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American Jf em Journal 


Published by the 








VOLUME 60 - 6 ( 


Missouri Botanicai. 
Gardes LiBfiAiiv 


Volume 60, Number 1, Pages 1-48, Issued April 21, 1970 

Aberrant Leaves on Angle-Shoots of Selaginella martensii Spring 

Terry R. Webster 1 

A Further Note on the Type of Platycerium alcicorne C. V, Morton 7 

Gametophyte Ontogeny and Sex Expression in Dryopteris ludoviciana 

Michael L Cousens and Harry T. Horner 13 

Shorter Nutes: A New Form of Microlepia speluncae; Two New State 
Records for Isoetes in Kentucky; Psilotum nudum New to Arizona. . . 28 

Notes and News 

Recent Fern Literature 


American Fern Society: Report of the President ; Report of the Secretary; 
Report of the Treasurer; Report of the Auditing Committee; Report 
of the Judge of Elections; Report of the Fern Spore Exchange 40 

Volume 60, Number 2, Pages 49-88, Issued July 3, 1970 

The Fern Collections in Some European Herbaria, VI C. V, Morton 49 

Fertile Sporophytes of Botrychium virginianum Attached to Gameto- '- 
pVtes Bruce W. McAlpin 62 

A Pecuhar Species of Grammitis C. V, Morion 65 

Vascular Organization of the Rhizome of Cibotium barometz 

Subhash Chandra 68 

A Survey of Some Morphological Features of the Genus Elaphoglossum 
in Costa Rica Robert M, Lloyd 73 

Notes and News , . . 83 



Recent Fern Literatiu-e 84 

American Fern Society yS 


Volume 60, Numher 3, Pages 89-128, Issued September 16, 1970 

Stelar Anatomy of Six Species of Lycopodium Frances M. Cardillo 89 

A Note on the Gametophytes of Cystopteris tenuiseeta 5. 5. Bir 98 




The Lectotype of Polypodium Icptophyllum L C. V. Morton 101 

Taxunomic Notes on Ferns, IV C. V. Morton 103 

A Reinvestigation of the ^Morphology of Hypodemailum crenatum 

B, K. Nayar and Nisha Bajpai 107 

Shorter Note: Pyrrosia princepts, a Fern New to Cultivation IIS 

Recent Fern Literature 119 

American Fern Society 128 

Volume 60, Number 4, Pages 129-172, Issued Decemher 18, 1970 

gA Major North American Range Extension for the Forked Spleenwort, 

■ Asplenium septentrionale David L. Emory 129 


Hard Water as a Limiting Factor in the Distribution of Isoetes echino- 

spora LeRoy Lee and Frances Belknap 134 

Indian Pteridophytes Used in Folk Remedies . . .H, S. Puri 137 

The Rhizome Scales of Platycerium Barbara Joe Hoshizaki 144 

Shorter Note^: Stenrf^Cross-section Prints Aid in Identifying Horsetails; 
Psilotum nudum Spreading Northward; Terminology of the Spor- 

angial Structures of Equisetum; Kinds of Cystopteris in California. . 161 

Notes and News ; 165 

Recent Fern Literature - 167 

American Fern Society ■ 169 

Index to Volume 60 1^0 

Errata 172 


January-Mahch, 1970 

No. 1 





Published by the 










MAY 1 1 1970 


fa^ROfiM LmnABx 

Aberrant Leaves on Angle-Shoots of Sclaginella martenBii Spring 

Tbbbt R. Websteb 

A Further Note on the Type of Platycerium alcicome . ^ „C. V. Morton 
Gametophyte Ontogeny and Sex Expression in Dryopteris ludoviciana 

Form of Microlepia speluncae 

Psilotum nudum New to Arizona 

Notes and News 

Recent Fern Literature ^. . . 

*-♦ •— • •-• 

9-9 9 



President : Report of the Secretary 

Report of the Treasurer ; Report of the Auditing Committee 
the Tudee of Elections: Reoort of the Fern Spore Exchange 



Michael I. Cousens and Harry T. Horner 13 




Elje American Jfern ^otietp 

Council foe 1070 

Warren H. Wagner, Jr., Department of Botany, University of Michigan, 
Ann Arbor, Michigan 48104. President 

John T. Mickel, New York Botanical Garden, Bronx Park, Bronx, New 
York. 10458. Vice-President 

A. Murray Evans, Department of Botany, University of Tennessee, Knox* 
ville, Tenn. 37916. Secretary 

LeRoy K. Henry, Division of Plants, Carnegie Museum, Pittsburgh, Penn- 
sylvania 15213. Treasurer 

David B. Lelunger, Smithsonian Institution, Washington, D. C. 20560. 


Rational ^ocietp 36itpxtientati\}tsi 

Warren H. Wagner, Jr., University of Michigan A,A.A.S. Representative 
RoLLA M. Tryon, Jr., Harvard University AJ.B.S. Representative 

American jfttn Sonrnal 


David B. Lellinger Smithsonian Institution, Washington, D. C. 20560. 

C. V. Morton Smithsonian Institution, Washington, D. C. 20560. 

RoLLA M. Tryon, Jr. 

Gray Herbarium, Harvard University, Cambridge, Mass. 02138. 

Ira L. Wiggins Dudley Herbarium, Stanford University, Stanford, Calif. 


An illustrated quarterly devoted to the general study of ferns, owned by 
the American Fern Society, and published at 3110 Elm Ave., Baltimore, 
Md. 21211. Second-class postage paid at Baltimore. The pages of the Jour- 
nal are open to members who wish to arrange exchanges; membership lists 
assist those interested in obtaining specimens from different localities. 

Matter for publication should be addressed to the Editor-in-Chief. 

Changes of address, applications for membership, subscriptions, orders 
for back numbers shoiud be addressed to the Treasurer, 

Subscriptions $5.50 gross, $5.00 net (agency fee $0.50); sent free to mena- 
bers of the American Fern Society (annual dues, $5.00; sustaining membership, 
$10.00; life membership, $100.00). Extracted reprints, if ordeml in advance, 
will be furnished authors at cost, plus postage. 

Back volumes $5.00 to $6.25 each; single back numbers of 64 pages or 

less, $1.25; 65-80 pages, $2.00 each; over 80 pages, $2.50 each; Cumulative 

Index to Volumes 1-25, 50 cents. Ten percent discount on orders of six volumes 
or more. 

Hflirarp anti J^eihatium 

Dr. W. H. Wagner, Jr., Department of Botany, University of Michigan, 
Ann Arbor, Michigan 48104, is Librarian and Curator. Members may borrow 
books and specimens at any time, the borrower p&ying all shipping costo, 

S>pore Cxtfjange 

Mr. Neill D. Hall, 1225 Northeast 95th Street, Seattle, Washington 98115, 
is Director. Spores exchanged and collection lists sent on request. 

^ifti anb 9tqntiti 

Gifts and bequests to the Society enable it to expand its services to mem- 
bers and to others interested in ferns. Herbarium specimens, botanicaJ books, 
back issues of the Joitrnal, and cash or other gifts are always welcomed, ana 
ve tax-deductible. Inquiries should be addressed to the Secretary. 

American Jfern S^ournat 

Vol. 60 January-March, 1970 No. 1 

Aberrant Leaves on Angle-Shoots of 
Selaginella martensii Spring^ 

Terry R. Webster 

Dorsal and ventral angle-meristems, which normally develop 
into roots (Webster and Steeves, 1967), are present at most points 
of shoot branching in the heterophyllous species Selaginella 
martensii Spring. But in stem segments that are surgically isolated 
from the intact plant and which are incubated on moist filter 
paper, these angle-meristems may develop as angle-shoots 
(Williams, 1937; Webster, 1969) {Fig, 6). In a recent study of 
angle-meristem development in excised stem segments of 8. 

7nartensii (Webster 

on angle- 

shoots were normal, according to the description by Harvey- 
Gibson (1897). But some, particularly those near the base of angle- 
shoots, w^ere abnormal. The morphology and possible significance 
of these aberrant leaves is presented in this paper. 

Normally, the larger, ventral leaves of the angle -shoots are 
lanceolate, w-ith a narrow, bluntly pointed apex {Fig. 1), The 


awn {Fig. 2), 

Both dorsal and ventral leaves possess marginal teeth and are 
asymmetrical at the base. A single median vascular strand is 
present, and a ligule occurs at the leaf base on the adaxial surface. 
Conspicuous rows of stomata occur along the vascular strand 
{¥ig. 7). 

^This research was supported by National Science Foundation Grant 
GB-7068 and by a grant from The University of Connecticut Research 


Volume 59, No. 4, of the Journal, pp. 137-176, was issued Jan. 14, 1970. 

American Fern Journal 

Volume 60, Plate 1 

-^^- . , *H- -^^W^■-■■'^^ ►^^VA 


r-m r-i J---r^^ M^*^^- r ^r ■ t -^rf r w r 

"A ■ ■ " ^ ■r'^ 







































Cleared and Stained Leaves of Selaginella martensu 

Angle -SHOOTS Sho\ving Vascular Strands 

Aberrant Selaginella Leaves 3 

In contrast to most leaves on angle-shoots, the first few leaves 
toward the base, proximal to the first branching of the angle- 
shoot, are aberrant {Figs, 3, 4, and 6), In addition to these abnormal 
basal leaves, a few cases of unusual leaves were found more distally 
on angle-shoots. Both dorsal and ventral aberrant leaves vary in 
shape. Instead of being entire, the leaves are variously lobed or 
divided at the tip (Figs, 3 and 4)- Ii^ some cases they are deeply 
cleft. Unlike normal leaves, the base of aberrant leaves is often 

For anatomical study, angle-shoots were cleared in 2 % sodium 
hydroxide and were stained with basic fuchsin according to the 
method described by Jacobs (1952). Instead of the normal single 
vascular strand, several strands occur in aberrant leaves {Figs. 3 
and 4)' As many as seven strands were observed in a single leaf. 
In most leaves one strand could be traced from the stem vascular 
system toward the leaf base. Near the base of the leaf the strand 
is divided, usually dichotomously, one to several times. A few^ 
ventral leaves were observed in which two separate strands could 
be traced from the stem into the leaf base. In neither normal nor 
aberrant leaves is the departure of leaf traces associated with a gap 
in the stem vascular cylinder. 

In normal and in most aberrant leaves the vascular system is 
closely associated with a single ligule. However, in some deeply 
cleft leaves, two separate ligules occur {Fig. 6) and one or more 
vascular strands are associated with each ligule. Ball (1925) 
reported sporophylls of *S. martensii which possessed two ligules, 
one of normal size and an additional smaller ligule. Double ligules 
found in the present study were of normal size {Fig. 5). In aberrant 
leaves each vascular strand has stomata associated with it, and, 
compared to normal leaves, more of the leaf surface is covered by 
stomata {Figs. 7 and 8). The arrangement of stomata in distinct 
rows is not so evident as in normal leaves. 

Fig. 1. Normal ventral leaf, X 23. Fig. 2. Normal dorsal leaf, X 22. 
Fig. 3. Aberrant ventral leaf, X 46. Fig. 4. Aberrant dorsal leaf, X 42. 
Fig. 5. Aberrant dorsal leaf with two ligules. Some stem tissue visible 
through leaf, X 47. 

American Fern Journal 

Volume 60, Plate 2 

Plate 2. Fig. 6. Diagram of ventral view of stem segment bearing 


vasculature toward base of angle-shoot. Fig. 7. Stomata on normal 


shoot. The abbreviations are: L = ligule. S =» stoma, V = vascular 


Aberrant Selaginella Leaves 5 


Grambast and Rosello (1965) described leaves of S. inlldenovii 
(Desv.) Baker in which there were two or three veins from the 
dichotomous branching of a single vascular trace. According to 
their illustrations, the leaves themselves were not so distinctly 
lobed or divided as the aberrant leaves described here in S. 
martensii. The authors made no mention of stomata or ligules in 
the leaves of *S. willdenovii, but they did discuss possible phylo- 
genetic implications of the branched vasculature. They cited 
fossil lycopods {ProtolepidodeMron Krejci, Duisbergia Kraus. 
and WeyL, and Sigillaria scutellata Brongn.) which possessed 
leaves with dichotomously branched vascular strands, and sug- 
gested that in S. ivilldenovn microphylls with dichotomizing veins 
represent a primitive condition. According to their view, micro- 
phylls with a single vein could have been derived by reduction. 



One could apply a similar interpretation to the microphylls 
of S. martensii. Aberrant leaves with branched vasculature could 
be interpreted as representing a primitive condition, and normal 
leaves with a single vascular strand a more advanced condition, 
A comparison of normal and aberrant leaves would suggest that 
during evolution lobing, number of vascular strands, and rows 
of stomata w^ere reduced. 

According to the theory of recapitulation (ontogeny, in 
abbreviated fashion, repeats phylogeny), the position of leaves on 
angle-shoots could be used to indicate a primitive or an advanced 
condition. Although usually applied to progressive development 
from juvenile to adult leaves in sporelings or seedlings, Takhtajan 
(1959, p. 91) has applied this theory to leaves occurring on acces- 
sory shoots. Angle-shoots would fit into this latter category. 
According to the theory, the progression of microphylls in angle- 



of angle-shoots would represent a primitive condition, and more 
distal (normal) leaves an advanced condition {Fig. 6). However, 

6 American Fern Journal 

as Foster and Gifford (1959; p. 9) have stated, whether juvenile 
leaves provide reliable evidence to the morphology of ancestral 
leaf types is questionable, and hypotheses based on the theory 
of recapitulation should always be made with great caution. It 
should also be noted that the features found in aberrant leaves of 
S. martensii are not known in fossil representatives of Selaginella 
or in Selaginellites, At the present time there is no strong evidence 
to indicate that aberrant microphylls in Selaginella do in fact 
represent a primitive condition. Rather than having any phylo- 
genetic significance, the aberrant leaves in S. martensii merely may 
be the result of abnormal development during the early ontogeny 
of angle-shoots. 

Literature Cited 

Ball, N. G. 1925. On the occurrence of a double ligule in Selaginella martensii. 

Ann. Bet. 39 : 884. 
Foster, A. S. and Gifford, E. M. 1959. Comparative Morphology of Vascular 

Plants. W. H. Freeman, San Francisco. 
Grambast, L. and S. Rosello. 1965. Les notions de microphylle et de mega- 

phylle et leur signification phylogenetique. Compt. Rend. Acad. Sci., 

Paris 261: 5183-5186. 
Harvey-Gibson, R. J. 1897. Contributions towards a knowledge of the 

anatomy of the genus Selaginella Spr. III. The leaf. Ann. Bot. 11: 

Jacobs, W, P. 1952. The role of auxin in differentiation of xylem around a 

wound. Amer. J. Bot. 39 : 301-309. 
Takht.uan, a. L. 1959. Essays on the evolutionary morphology of plants. 

(Transl. by O. H. Gankin) Amer. Inst. Biol. Sci., Washington, D.C. 

Webster, T. R, 1969. An investigation of angle-meristem development in 

excised stem segments of Selaginella martensii. Canad. J. Bot. 47: 

and T. a. Steeves. 1967. Developmental morphology of the 

root of Selaginella martensii Spring. Canad. J. Bot. 45 : 395-404. 
Williams, S. 1937. Correlation phenomena and hormones in Selaginella. 

Nature 139: 966. 

Systematic and Environmental Biology, University of 
Connecticut, Storrs, Conn. 06268. 

TyrE OF Platycerium alcicorxe 7 

A Further Note on the Type of Platycerium alcicorne 

C. V. Morton 

Several years 'ago I published a paper, 'The Nomenclature of 
a iMadagaBcarian PlatyceriijSii'' (Morton, 19G4), in which I con- 
cluded that A c^ostichum alcicorne Swartz (Syn. Fil. 12, 196, 180G) 
was an iliegitimale change of name for Acrostichum stemaria 
Palisot (Fl. d'Oware et Benin 3: t. 2. 1805), which was cited by 
Swartz as a synonym. This was an error on my part, occasioned by 
the fact that nowhere in the "Index Filicum" or in Willdenow or 
Desvaux is there any indication that Swartz had published his 
Acrostichum alcicorne prior to 1806. Mr, De Joncheere was the first 
to note that the name A. alcicorne Swartz was actually published 
in Schrader's "Journal fiir die Botanik,'* (ISOO, vol. 2, part 1, p. 
11, 1802).^ Since this 1802 publication is prior to the publication of 
A, stemaria Palisot it is clear that A. alcicorne Swartz was not a 
renaming of A, stemaria, and must therefore have a different type. 

Swartz' 1802 paper "Genera et Species Filicum" is a sort of 
preliminary to his extensive "Synopsis Fihcum" of 1806; it is very 
brief throughout, with short diagnoses and no citation of speci- 
mens, localities, or synonymy in general. In the case of A, alcicorne 
Swartz did cite '^Neuroplatyceron Pluk. am. phyt. t, 4^9yf. 2J^ De 
Joncheere (1967) has argued that this Plukenet plate must be 
taken as the lectotype of -A. alcicorne. However, it was not Swartz^ 
habit to give names indiscriminately, if ever, to plants known to 
him only from pre-Linnaean drawings. In this case there is ample 
evidence that the protologue included not only this pre-Linnaean 
plate of Plukenet but also herbarium material y for Swartz described 
the fertile fronds, which were not depicted by Plukenet. In cases 
like this there is general agreement that species ought to be 
typified by specimens when possible and not by plates. 

^ The date ''1801'^ on the title page has usually been accepted, but Staflea 
in his "Taxonomic Literature/' p. 432. 1967, has shown that this part was 
pubHshed in March, 1802. 

Amehican Fern Journal 

Volume 60, Plate 3 

- --/-,- *r-- 

_ - - ^^^ * -KJ "A-" J ■ n ■ r ■- M_^ ■ f|A %0 > -ii_-J -ii_-_--->- -'-■'j^j r'r,^^ 


A*iV---"-".^ ^*^ ^fit 






U,.j — L™ 

Museum bolanicum Up&aiiense 


^ / 



. ^ 

'y^ ' ^ n^\ ^ 

^ > .^MViW --^i 



Lectotypevof Acrostichum alcicorne, nest leaves (UPS) 

Type of Platycerium alcicorne 9 

Dr. Schelpe searched the herbarium in Stockhohn and could not 
find a specimen suitable for a lectotypc, and De Joncheere stated 
that Schelpe also searched the herbarium in Uppsala without 
finding any specimens. At my request, Dr. John Wurdack re- 
cently found that there are indeed specimens in Uppsala, and 
through the courtesy of Dr. Rolf Santesson I now have these on 
loan. The specimens mentioned are in the original herbarium of 
Afzelius. They are fine specimens, including nest-leaves and fertile 
leaves, mounted on five sheets. They are indicated as being from 
Sierra Leone collected by Smeathman. In his ISOG publication 
Swartz mentions (p. 196) collections from Madagascar, Commerson; 
Oware, Palisot; Java, Thunherg\ Guinea, Afzelius; and Xova 
Hollandia, Xee," and on p. 12 cites ''Sierra Leone" as one of the 
localities. In this 1S06 publication Swartz had added as syno- 
nyms A, stemaria Palisot and A, hifurcahim Cav., and it is likely 
that the collections of Commerson and Palisot were cited from 
Palisot rather than actually seen by Swartz and that the Nee 
collection was cited from Cavanilles. No specimen from Java 
collected by Thunberg has been found. The Afzelius collection 
from Sierra Leone mentioned above is the only herbarium speci- 
men that can definitely be associated with a collection cited by 
Swartz. The name alcicorne is in the hand of Swartz. The five 
sheets in the Uppsala Herbarium are here designated as the 

lectotype of A. alcicorne Swartz. According to Schelpe, there is a 

specimen in Stockholm from Sierra Leone annotated by Wikstrom 

as A. alciconiey and this is doubtless an isolectotype, although 

Afzelius is not definitely indicated as the source. 

The specimens from Sierra Leone designated as lectotype are 
identical with A. stemaria Palisot from Oware [Nigeria], the type 
of which I have seen in Paris. The Plukenet plate was apparently 
drawn from a specimen now in the Sloane Herbarium in the British 
jMuseum, and according to De Joncheere came from the Comoro 
Islands and is identical with the plant described as Acrostichum 
alcicorne Willemet, now properly Platycerium vassei Poisson. 
The lectotype chosen above agrees well enough with Swartz' 

American Fern Journai. 

Volume 60, Pl 



■ d I »■ T r F T_V 















WuseurtT botanicum UpsaUense 



J.. r 

!t!^u MxetSitfk. 

/^'-A^/'A ' -* .» 

» Tyrt »,W,,, *'^>„*,^^ , ^ 


Type of Platycehium alcicorne 11 

brief diagnosis, at least as well as that diagnosis agrees with P. 

The present lectotypification is really best for stability as well as 
fully justified by the facts. Although A, alcicorne Swartz was 
prior to A, stemaria Palisot by three years, it was itself illegiti- 
mate, being a later homonym of A. alcicorne Willemet (179G), 
a species based on an entirely different type (Madagascar, Sladl- 
mann). The next earliest name for the West African species is 
thus A, stemaria Palisot, and the correct name for it is Platyceriuni 
stemaria (Palisot) Desv., of which P. alcicorne (Swartz) Desv. is 
thus a taxonomic synonym. This is a fortunate turn of events, 
since P. stemaria is a name well known among horticulturists. 
Incidentally, Dr. Schelpe has suggested in a letter to Barbara 
Joe that the epithet ' 'stemaria" ought to be corrected to 
stemmaria," since the root is from '^stemma," a garland, in 
Greek. This may well be, but the correction of the original spelling 
of a name ought not to be made for purely etymological reasons if 
there is no reason to suspect a purely typographical error; there are 
many poorly or wrongly formed names that must nevertheless be 


The name P. alcicorne Desv. adopted by De Joncheere for the 
East African and Aladagascarian species is thus incorrect, which is 
fortunate, since this epithet has been widely misapplied to three 
different species — the West African P. stemaria^ and the East 
African and ^ladagasearian P. vassei, and the Australian P. 

Another species of Plafycerium that may be mentioned here 
incidentally is P. anyolense Welw. ex Baker in Hook & Bak. Syn. 
Fil. 425. 1868. The name appears as a comment under P. aethiopi- 
cum Hook, as follows: '^Dr. Welwitsch has furnished us with a 
plant (P. angolense Welw. MSS.) which is most like this, but has 
a broad-cuneate fertile frond 9 in. br. at the top, without either 
forks or horns, and with a patch of fruit nearly as broad as the 
lamina." Dr. Schelpe has suggested in a letter to me that this name 
ought to be rejected by Art. 34 of the Code as not being accepted 
by the publishing author. In my opinion it is valid, as valid as a 

12 American Fern Journal 

large number of other names that are published rather casually 
in footnotes or text-notes. Baker did not either definitely accept it 
or reject it. He certainly did not conclude that it was exactly the 
same as P. aethiopicunij but gave its differences as he saw them. 
Since there is a name and a description, the name ought to be 
accepted as validly published. As it turns out, Welwitsch was 
quite right that it was different from P, aethiopicum Hooker, 
and P. angolense Welw. ex Baker is the correct name for the third 
African species of Platycerium, the other two being Platycerium 
stemaria and P. vasseij a taxonomic synonym of P. angolense is 
P. elephantotis Schweinf. and another probable synonym is P. 
velutinum C. Chr., as discussed by Barbara Joe (1964) in her 

useful review of Platycerium. 

Literature Cited 

Joe, Barbara. 1964. A review of the species of Platycerium (Polypodiaceae). 

Baileya 12 : 69-126. 

Joncheere, G. J. DE. 1967. Notes on Platycerium Desv. I. Nomenclature and 

typification of the genus and species in Desvaux's original publica- 
tion of 1827. Blumea 15 : 441-451. 

Morton, C. V. 1964. The nomenclature of a Madagascarian Platycerium. 

Baileya 12: 36-38. 

National Museum of Natural History, Washington, D. C. 

Dryopteris Gametophytes 13 

Gametophjrte Ontogeny and Sex Expression in 

Dryopteris ludoviciana^ 


Although the ontogeny of fern gametophytes has a long history 
of morphological investigation, sex expression in ferns has only 

recently been studied by experimental morphologists and geneti- 
cists. ]\Iost homosporous ferns have bisexual gametophytes whose 
mitotically derived gametes produce a completely homozygous 
sporophyte after selfing (Klekowski and Baker, 1966). Wilkie's 
work (1956), which demonstrated self incompatibility in Ptericlium 
aquilinum, has stimulated the use of fern gametophytes in genetic 
research. lOekowski and his associates (1968, 1969) used genetic 
and morphological criteria to characterize pteridophyte gameto- 
phytes in terms of their probability for intragametophytic (func- 
tionally bisexual) and intergametophytic (functionally unisexual) 

This study w^as undertaken to determine whether the ontogeny 
of Dryopteris ludoviciana (Kunze) Small gametophytes differed 
from that reported for other dryopterid and thelypterid gameto- 
phytes (Kny, 1895; Waldemann, 1928; Xayar and Chandra, 1963), 
and to determine sex expression under controlled conditions. 
The effect of population density on sex expression was tested by 
culturing single isolated gametophytes and gametophytes in moder- 
ate and dense populations. The ontogeny and sex expression of 
Thelypteris denlala (Forsk.) E, St. John was also observed for 

Methods and Materi.\ls 

Dryopteris ludoviciana occurs in cypress and gum swamps, 
as well as in lune sinks, from Florida west along the Gulf Coast to 
Louisiana and north as far as North Carolina (Brown and Correll, 

1 This paper is a portion of a thesis completed by the senior author for the 
Master of Science Degree, I. S. U., 1969. 

14 American Fern Journal 

1942). A detailed record of its occurrence is not available^ but it 
is less common than other Dryopteris species. 

Two Dryopteris ludopiciana sporophytes were greenhouse- 
grown after their collection on November 20, 1966^ in Alachua 
County, Florida, in the Devil's Alillhopper, a lime sink 5-6 miles 
northwest of downtown Gainesville. A voucher specimen {John T, 
Michel 1730) is in the Iowa State University Herbarium. The 
plants used in this study were diploid (2n = 82) (A. R. Smith, 
pers. comm.)- 

Fertile pinnae were cut from vigorous fronds and rinsed in 
running distilled water to remove foreign spores. The pinnae 
were then sealed in small manila envelopes. After most of the 
sporangia had dehisced (8-12 days at 20°C) the spores were trans- 
ferred to small corked vials and utilized within two months. 

All spores were sown on a steiile mixture of loam and coarse 
sand (4:1). Varying numbers of spores were sown to produce 
different densities. Population density was determined by counting 
all gametophytes visible at 30 X. The average number of gameto- 
phytes per cm^ was then determined. Dense populations contained 
10-21 gametophytes per cm^ and moderate populations had 1-3 
per cm^. Isolated spores were cultured in small petri dishes. 

Culture vessels containing only sterile soil were opened for 
watering and observation with the same frequency as the experi- 
mental cultures. Fern gametophytes appearing in these cultures 
reflected the extent of contamination (about one per cent). Densely 
sown cultures of Thelypteris dentata were also maintained under the 
same conditions of culture as Dryopteris ludoviciana and later were 
surveyed for the occurrence of gametangia. 

Conditions of culture, including the use of soil as a substrate 
(Atkinson and Stokey, 1964), were selected to approximate the 
habitat of naturally occurring fern gametophytes. The cultures 
were maintained in a Percival growth chamber. Diffuse illumina- 
tion was provided by white fluorescent tubes and incandescent 
bulbs at an intensity of 350-400 ft-c. Temperature at the soil 
surface was maintained at 22°C during the 12 hr light and 12 hr 
dark periods. 

Dryoptekis Gametophytes 15 

Samples of 24 or more gametophytes were removed for 
observation of development and sex expression at 25, 35, 45, 58, 
72, and 111 days after sowing. An effort was made to select gameto- 
phytes representative of all sizes and stages. The gametophytes 
were placed in a petri dish or on a glass slide containing 1.75% 
lactose solution for microscopic examination. This medium pre- 
vented plasmolysis for 8-12 hours and did not hinder the dehis- 
cence of antheridia or the motility of sperms. 

Gametophyte Ontogeny and Gametangia 

Spores of Dryopteris hidcwiciana are bilateral with a prominent 
perispore. A spring-collected sample (April to June) of 162 spores 
ranged from 24-50 /x long by 16-33 ^ wide. One hundred and eight 
fall-collected spores (October) ranged from 21-54 /x long by 16-37 
M wide. The spring-collected spores averaged 36 X 24 /x and the 
fall-collected spores averaged 36 X 26 /x. Spore size was con- 
siderably less uniform in the fall collection {Fig. i), but the bio- 
logical significance of this^ if any, is not known. A bimodal dis- 
tribution of spore size was not found. 

Most spores germinated within 6-20 days, but some took as 
long as three months. The spore coat ruptured irregularly or along 
the ridge and remained attached to the basal cell (Fig, 2), The 
first rhizoid was delimited shortly after germination, and a second 
rhizoid sometimes formed before filamentous growth began {Fig. 
3). Successive divisions then produced a 4-6-celled filament which 
was either stout {Fig, 4) or elongate {Fig. 5). Filamentous growth 
was prolonged if a spore germinated in a soil crevice, or was 
otherwise shaded {Fig. 7). These attenuated filaments eventually 
became planate. 

Longitudinal or oblique divisions in the terminal cell of the 
filament initiated planate growth {Fig. 6), These divisions could 
occur when the filament was only two or three cells long {Fig. 8). 
Subsequent longitudinal divisions could occur in all but the basal 
cell of the filament, obscuring the earlier filamentous form {Fig, 9). 

Continued divisions in two dimensions were most frequent 
distallv, Droducine: a trianeular eametonhvte of 10-15 cells with 

16 American Fern Journal 

marginal trichomes (Figs. 11, 12), This stage was occasionally 
irregular {Fig. 10), but these irregularities Avere obscured by sub- 
sequent symmetrical growth. The fastest growing gametophytes 
estabhshed an apical cell 12-16 days after germination {Fig, 13; 






























• • ••^•/^ oo ^# o o 




o o 

18 I O 

I6| • OO 

20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 30 52 54 


Fig. 1. Size Distribution of Dryopteris ludoviciana spores. O = one 


Table /). Slower growing gametophytes reached this stage within 
16-30 days. The apical cell cut off derivatives laterally to produce 



were rare, and all gametophytes, except ameristic males which 
lack a notch meristem, became cordate {Fig. 16). 

Trichomes were common on the dorsal surface, averaging one 
trichome per five prothaUial cells. This ratio diminished to one 

Dryopteris Gametophytes 17 

trichome per fifteen prothallial cells on the wings and to one per 
20-30 prothallial cells on the ventral surface. 

Gametophytes 20-45 days old and 0.5-0.7 mm wide initiated a 
cushion which \vas two or three cells thick at first and later four 
or five cells thick. Lateral prothallial cells {Fig, 20) became much 
larger than younger anterior cells {Fig, 19), Older gametophytes 
were 0.9-13.0 mm wide {Figs, 17 y 21), depending on their pattern 
of sex expression. 

Table I. Initiation (in days after germination) of the Stages of 

Development of Dryopteris lvdoviciana gametophytes. 

Rapidly growing Slow growing 

Developmental stage gametophyte ganieiophyle 

Filamentous 6 6 

First longitudinal division 7 7 

Early planate (9-15 cells) 8-11 11-29 

Apical cell 

12-16 17-30 

Notch meristem 14-19 20-32 

Cushion initiation 20-26 27 

- J ai 

The antheridia are of the polypodioid type (Atkinson and 
Stokey, 1964), consisting of a single basal cell, a cylindrical 
middle cell which surrounds the central primary androgonial cell, 
and an undivided cap cell. 

The archegonia are of the advanced type (Atkinson and 
Stokey, 1964) with four or five tiers of neck cells, each tier con- 
taining four cells. The central canal cell was binucleate and en- 
larged distally. The archegonium w^as slightly larger at its apex 
than at its base and was always inclined slightly toward the base 
of the gametophyte- Archegonia occurred ventrally on a broad 
area just behind the notch, and up to SO occurred on gameto- 
phytes 8-13 mm wide. 

Sex Expression in Popul.a-TIOns 

Dryopteris ludoviciana has a normal sexual alternation of 
generations. The gametophytes observed in isolated, moderate. 

American Fern Journal 

Volume 60, Plate 5 

Plate 5. Early Ontogeny op D. ludoviciana Gametophytes, The 



Figs. 7, 15; 1.5 cm 

100 m fob 



DKYorTERis Gametophytes 19 

and dense cultures expressed one of five possible patterns of 
sexuality, or lacked gamctangia. The five sex expression patterns 
were based on the presence of one or both types of garnet angia, 
and, where both kinds of gamctangia occurred, the sequence of 
their appearance. The presence or absence of a notch meristem 
was also recorded for all gametophytes. 

Rapidly growing gametophytes {Table I) usually became uni- 
sexual females and were the first plants in all populations to become 
sexual {Fig. 18). Archegonial initials were found on 24-day-old 
gametophytes as small as 0.9 mm wide. These gametophytes 
continued to grow and produce additional archegonia, attaining 
an average width of 4.6 nun 45 days after germination {Fig, 21), 
These gametophytes remained unisexual after 105 days and 
became 8-13 mm wide {Fig, 17). Older gametophytes occasionally 
produced basal outgrowths which became antheridial. 

Protandrotis bisexual gametophytes are tjT3ical of most homo- 
sporous leptosporangiate fems (Atkinson and Stokey, 1964), but 
were rare in D. ludoviciana populations studied here. These 
gametophytes first produced antheridia among the rhizoids and 
on the basal half of the cushion. Six to 20 days later archegonia 
formed below the notch meristem. Antheridia numbered fewer 
than 80 on protandrous bisexual gametophytes. Twenty of the 
527 gametophytes surveyed were protandrous bisexual. 

In synchronous bisexual gametophytes archegonia occurred in the 
typical position below the notch meristem and antheridia were 
restricted to the distal half of the cushion adjacent to and among 
the archegonia. ^Maturation of both kinds of gametangia was 
S3aichronous, so that the periods of antheridial dehiscence and 
archegonial receptivity overlapped. These gametophytes first 
became sexual 30-40 days after germination and continued to 
produce both kinds of gametangia. After 45 days a gametophji;e 


bore an average of ten archegonia and 22 antheridia. 

Ameristic male gametophytes developed antheridia precociously 
but failed to develop an organizing notch meristem. They became 
sexual shortly after germination, producing antheridia on almost 
every cell, and occasionally two per cell. Ameristic males devel- 

American Fern Journal 

Volume 60. Plate 6 

1 [Jim 


Late Ontogeny of D. ludovicl\na Gametophytes. The 

numbers in circles indicate days after germination. flg. 
growing gametophyte with cushion but without gametangia 
Unisexual female gametophyte with overlapping wings ani 
notch; abchegonia not illustrated. Fig. 18. RAProLY growing 
phyte with archegonia. Fig. 19. Cells from anterior margin 
OF. Fig. 20. Fig. 20. Celi^ from lateral area of Fig. 21. Fio. 21. 
growing unisexual female gametophyte. 

Dryopteris Gametophytes 


oped only from spores which either remained within sporangia, 
were next to sporangial fragments, or were in contact with the 
rhizoids of older gametophytes. Their form varied from filamentous 
to spathulate. 










Per cent 



of total 


size mm 

25 35 45 

25 35 45 

25 35 45 

- 75 

- 12 

- 13 

100 - 100 

34 - 6 I 1.9 - 2.6 





2.1 - 3.6 

90 - 1 I 0.4 - 4.0 

126 - 8 


, Per cent 
of totdl 
25 35 45 



25 35 45 


size mn 

25 35 45 

43 60 


13 la 


1.6 3.3 6.1 






- 3.9 6,3 

- . - 

_ - _ 

57 17 


17 5 


0.4 0.5 0.8 

100 100 


30 30 


_ - _ 



34 40 39 I 10 14 11 


- - 

1.5 2.9 3.7 






7 28 36 

2 10 10 


3 10 

1.8 2.5 3.1 

48 30 25 I 15 II 7 

100 100 100 I 30 36 28 

0.6 1.4 - 

0.5 0.3 0.9 








43 - 30 I 21 - 


- 2 

- 1? 

- 36 

- 2 

5.3 - 3.6 

- 3.5 


7 - 9 I 0.8 - 0.6 

^ 20 I 16 - 14 I 1.7 - 1.5 

- 26 I 2.0 - 2.5 

100 - 100 I 49 - 73 

40 30 32 I 29 27 26 

8 15 22 

32 27 


6 13 18 

20 28 36 I 14 25 29 

23 24 7 

100 100 100 I 72 89 81 

2.4 3.0 3.5 

- 5.0 

2.7 3.1 3.y 

0.8 0.6 0.9 

0.6 0.6 0.5 

Table II, Sex Expression in 25,35, and 45-d ay-old Isolated, Moderate, 
AND Dense Dryopteris litdoviciana Cultures. 

Meristic males produced 100-300 antheridia after they became 
cordate. The antheridia occurred on two-thirds of the wings, and 


American Fern Journal 

extended to within four cells of the notch meristem, Meristic males 
continued to produce antheridia after being transplanted back to 
soil, and were not bisexual after 105 days, 










? f 1^ ?■ •" fl' NO 

? f 4 !^ •' d" ••o 

3S DtIS 





S ^ if rf- 0* NO 

5 5 a;? ■• a N 


15 0»rS 


i i i.i ^ ^ HO 

2t HYS 



« ^ (f,f •■ <r MO 

4S 0«11 

49 Dili 

^ ^ j/f ^ d- HO 

4S Sirs 






Fig. 22. Average of Sex Expression for Spring and Fall Cultures 
Combined. The symbols represent unisexu.\l females, synchronous 


The occurrence of meristic males, amcristic males, and protan- 
drous bisexual gametophytes, and the frequency of all five sex- 
expression types, varied with population density {Table II). 
Neither type of male occurred in isolated cultures, whereas 
moderate cultures 45 days and younger contained two to ten per 

DRYorxERis Gametophytes 23 

cent meristic males. Dense cultures had both types of unisexual 
male gametophytes at a 28-33 percent level. 

Unisexual female and synchronous bisexual gametophytes 
occurred in all 25-day-old cultures {Table II; Fig. 22). Most of the 
unisexual females bore receptive archegonia at this time; most 
gamctangia on synchronous bisexual gametophytes were immature. 

Sexuality was expressed rapidly in dense cultures^ as only 10-23 
per cent of the gametoj^hytes in 25-day cultures were without sex 
organs. Forty-eight per cent of the gametophytes in 25-day moder- 
ate cultures were asexual, whereas isolated cultures were slowest, 
with 57-71 per cent remaining asexual at 25 days. The proportion 
of gametophytes without sex organs decreased with time in all 
cultures from an average of 19 per cent at 45 days {Table II; Fig. 
22) to zero at 105 days. 

Protandrous bisexual gametophytes were first found in dense 
cultures at 45 days and they remained at a two to four per cent 
level during 105 days of culture. The proportions of other sex ex- 
pression classes in dense cultures remained approximately uniform 
through time, except for the absence of ameristic males at 35 
days {Table II; Fig, 22), 

The proportion of synchronous bisexual gametophytes increased 
with time in moderate cultures, but the proportion of unisexual 
females remained uniform. In 58 and 72-day-old moderate cul- 
tures protandrous bisexual gametophytes occurred at a 10-15 
per cent level and meristic males increased to 31-40 per cent, ' 

The proportion of unisexual females in isolated cultures in- 
creased from 35 per cent at 25 days to 77 per cent at 45 days, 
while the proportion of unisexual females in moderate and dense 
cultures varied Uttle from the initial percentage. 

The Thelypteris dentata gametophytes which were cultured 
under the same conditions as Dryopteris hidoviciana expressed 
the protandrous bisexual pattern in all observed cases. 


The ontogeny of Dryopteris ludoviciana as described here does 
>t differ sienificantlv from the typical cordate pattern of the 

24 American Fern Journal 

Polypodiaceae sensu Into. The ontogeny of D. ludoviciana does 
not distinguish it from either Aspidium spinulosum Swartz 
[Dryopteris spimdosa (^luell) Watt.] or A. thelypteris Swartz 
[Thelypteris palustris Schott] as reported by Waidemann (1928). 
Kny's (1895) illustrations of the development of Dryopteris 
filix-mas gametophytes fail to distinguish it from D. spinulosa, D. 
ludoviciana, and Thelpyteris palustris. It is hkely that both Dryo- 
pteris and Thelypteris gametophytes have evolved to the same 
speciahzed level, and that gametophyte morphology is of little 
value in distinguishing between dryopterid and thelypterid ferns. 
The five sex expression types found in Dryopteris ludoviciana 
populations allow intragametophytic mating and two levels of 
intergametophytic mating. Intragametophytic fertilization elimi- 
nates heterozygosity. Intergametophytic fertilization between 
gametophytes from the same sporophyte is equivalent to self- 
pollination and reduces heterozygosity. Intergametophytic fertili- 
zation between gametophytes derived from different sporophytes 
is equivalent to cross-pollination and increases heterozygosity 
(Klekowski, 1969). 

The first sexual gametophytes observed in all populations were 
unisexual females, and the potential for intergametophytic fertili- 
zation was initially great. Intragametophytic fertiUzation became 
possible five to fifteen days later when s5aichronous bisexual 
gametophytes matured. 

Although the pattern of spore distribution for naturally occur- 
ring D. ludoviciana plants is not known, it is likely that a greater 
number of spores will fall near the parent sporophyte and that 
spore density will decrease with distance from the parent plant. 

Fertilization is most likely in dense populations because all five 
sex expression types occur and gametophytes often overlap. 
Fertilization is less likely in moderate populations and is least 
likely in isolation. 

Dense populations from a single sporophyte have a high likeli- 
hood of intergametophytic gamete exchange which reduces hetero- 
zygosity, but are unlikely to effect exchange with gametophytes 
derived from different parent sporophytes. Synchronous bisexual 

DiiYOPTERis Gametophytes 25 

gametophytes would maintain a low level of intragametophytic 
fertilization in dense populations (Table II; Fig, 22). ^Moderate 
populations have the highest level of intragametophytic fertiliza- 
tion because of the high proportion of synchronous bisexual 
gametophytes. The potential for intergametophytic fertilization is 
lower than that of dense populations because fewer gametophytes 
overlap. Decreased population density would, however, increase 
the likelihood of establishment of a foreign spore. This would in- 
crease the possibility of intergametophytic fertilization at a level 
which increases heterozygosity. 

Isolated bisexual gametophytes could establish sporophytes by 
selfing. Isolated unisexual female gametophytes would fail to es- 
tabhsh a sporophyte, except where spore distribution patterns 
of different D, ludoviciana plants overlapped. The resulting sporo- 
phyte would greatly increase heterozygosity in the gene pool. 

Variation of sex expression types found in different population 
densities imphes the existence of a population factor which in- 
fluences sex expression. The occurrence of unisexual males in dense 
and moderate cultures, but not in isolation, indicates that this 
factor influences maleness. Dopp (1950), who discovered an 
antheridium-inducing substance in Pteridium agnilimim, found 
that the Pteridium factor induced precocious antheridium forma- 
tion in Dryopteris filix-mas, and an aqueous extract from D. 
filix-mas gametophytes induced antheridia on P. aquilinum. 

Naf (1958, 1961) determined the mode of action of the Pteridium 
factor in P. aquilinum populations; his findings parallel certain 
aspects of the Dryopteris ludoviciana system. He found that uni- 
sexual females, ameristic males, and protandrous bisexual gameto- 
phytes occurred in P. aquilinum populations. Isolated gameto- 
phytes became unisexual females only, while all gametophytes 
became ameristic males if cultivated under conditions of slow 
growth with added Pteridium factor. Susceptibility to the induction 
of antheridia ceased before archegonia were formed, and the fastest 

growing gametophytes in the population probably attained the 
insensitive phase before the concentration of the factor was 
sufficient to induce antheridia. 

26 A:\iERicAN Fern Journal 

The fastest growing Dryopterts ludoviciana gametophytes also 
become unisexual females, and maleness does increase with popula- 
tion density. The occurrence of bisexual gametophytes in isolation, 
however, cannot be the result of an antheridium-inducing sub- 
stance secreted by other gametophytes. The antheridium-inducing 
factor also fails to explain the typical bisexual pattern of the 
majority of leptosporangiate ferns. 

It is possible that the typical bisexual condition of most fern 
gametophytes is the ancestral condition, and that the P. aquilinum 
system is derived. In this latter case all spores are potential 
unisexual female gametophytes, and other sex expression types are 
determined by the interaction of an antheridium-inducing sub- 
stance and variation in growth rate. If this is so, then the D. 
ludoviciana system is an intermediate one which shares the ad- 
vantages of both the ancestral and advanced conditions. 

At its present level of evolution the D. ludoviciana system has 
three favorable adaptations which increase its evolutionary po- 
tential. Sporophyte production is guaranteed in all population 
densities by a persistent level of the ancestral bisexual condition. 
Where a sporophyte population is already established; the level 
of heterozygosity is maintained by two levels of intergametophytic 
fertilization, so that the ability of the population to adapt to a 
changing environment is not lessened. The potential to increase 
heterozygosity is highest at the limit of the range so that range ex- 
tension is possible by gamete exchange between neighboring D. 
ludoviciana populations. Finally, unlike P. aquilinum, some 
isolated spores can produce bisexual gametophytes which may 
self to form a sporophyte. This would allow range extension of 
the species throughout a habitat and disjunct range extension 
when scattered spores occur in a suitable habitat far removed 
from that of the parent sporophyte. 

The role of the gametophyte in the natural distribution of ferns 

has not been assessed. Fern spores do act as long distance 
propagules, but the establishment of a gametophyte does not 
guarantee sporophyte occurrence (Farrar, 19C7). Both gameto- 
phyte and sporophyte are subjected to selection in range exten- 

Dryoptekis Ga.metophytes 27 

sion of the species. Gametophyte adaptations, \vhich would seem 
to encourage colonization, may be ineffective if the sporophyte 
is limiting. The relative effectiveness of gametophytes and sporo- 
phytes in colonization, the frequency of gametophyte establish- 
ment, and the occurrence of sex expression types must now be 
determined in the field. 

Literature Cited 

Atkinson, L. R., and A. G. Stokey. 1964. Comparative morphology of the 

gametophyte of homosporous ferns. Phytomoiphology 14: 51-70. 

Brown, C. A., and D. S. Correll. 1942. Ferns and Fern Allies of Louisiana. 

Louisiana State Univ. Press, Baton Rouge, Louisiana. 

Dopp, W. 1950. Eine die Antheridienbildung bei Farnen forderne Substanz in 

den ProthalHen von Pteridium aquilinum (L.) Kuhn. Ber. Deut. 

Bot. Ges. 63: 139-147, 


independently of their sporophytes in the Southern Appalachians. 
Science 155: 1266-1267. 
Klekowski, E J., Jr. 1969. Reproductive biology of the Pteridophyta. II. 

Theoretical considerations J. Linn, Soc. London (Bot.) 62 : 347-359. 

gnificance of polyploidy 

in the Pteridophyta. Science 153: 305-307. 

, and R. M. Lloyd. 1968. Reproductive biology of the Pterido- 
phyta 1. General considerations and a study of Onoclea sensibilis 
L. J. Linn. Soc. London (Bot.) 60: 315-324. 

Kny, L. 1895. Entwicklung von Aspidium fihx-mas. Bot. Wandtafeln 9: 


Naf, XJ. 1958. On the physiology of antheridium formation in the bracken 

fern (Pteridium aquilinum (L.) Kuhn), Physiol. Plant. 11: 728-746. 

. 1961. On the physiology of antheridium formation in ferns, pp. 

709-723. In R. M. Klein, ed. Plant Growth Regulation. Iowa State 
Univ. Press, Ames, Iowa. 

Nayar, B. K.j and P. Chandra. 1963. Observations on the morphology of 

the gametophyte of Cyclosorus. J. Indian Bot. Soc, B, 42: 392-400. 

Wat^demann, H, 1928, Beitrage zur Entwicklungsgeschichte der Prothallien 

einheimischerPolypodiaceen. Dissertation, Univ. ^larburg, Germany. 

WiLKiE, D. 1956. Incompatibility in bracken. Heredity 10: 247-256. 

Department of Botany and Plant Pathology, Iowa State 
University, Aaies, Iowa 50010. 

28 American Fern Journal 

Shorter Notes 

A New Form op Microlepia speluncae. — ^A cultivated form 
of Microlepia that is a very graceful and delightful plant for 
greenhouses (and possibly outdoors in southern California) has 
been introduced rather recently. It was in cultivation in the 
Botanical Garden of Ajuda, Lisbon, Portugal, and material was 
brought back by Dr. F. G. Meyer to the Department of Agri- 
culture, where it is in cultivation in Glenn Dale, Maryland (Plant 
Introduction no. 241,924). In Portugal it was passing as Micro- 
lepia strigosa (Thunb.) K. B. Presl cv, 'Cristata'; however, the 
plant is quite different from M. strigosa, which is a related species 
also in cultivation. In habit it is quite like the plumed form of the 
English Lady Fern, Athyrium filix-femina cv. ^Corijmhiferum^ 


form of M. sveluncae (L,) Moore 





bipimiate or bipinnate-pimiatifid. .M 

Veins (at least in dried material) not raised above the surface beneath; hairs 
of rhachises and midribs fine, horizontally spreading; blades tripinnate to 
quadripinnate M. speluncae 

Microlepia speluxcae cv. 'Corymbifera* Morton, cv. nov. 

Main rachis 4- or 5-tmies forked, forming a plumelike tip; 
lateral pinnae mostly drooping, forked and plumose at the tips. 

Type: Botanical Garden of Ajuda, Lisbon, Portugal, August 10, 
1957, F. (7. Meyer 5096 (US 2,292,318). 

This plant is also in cultivation at Longwood Gardens, Kennett 
Square, Pennsylvania (Ace. no. 58,498), grown from material 


Meyer, Wikoff 1200 (US 2,292.212-3), Huttlestm 


for I recall seeing simi 
Morton, U. S. National 


C. V. 

American Ferx, 

Volume 60, Plate 7 



''/ 4 ^ ^ 

<xxn»AtiD njwTft 

V ^ 

> ^ 


30 AiviERicAN Fern Journal 

Two New State Records for Isoetes in Kentucky. — The 
genus Isoetes has not been known positively for the state of Ken- 
tucky. Davies (Ann. Kentucky Nat, Hist. 1: 33-44. 1949) re- 
ported Isoetes engelmannii A. Braun in Jefferson County, but 
searches in the area in later years failed to confirm its existence 
there, according to Reed (Castanea 26: 94-96. 1961). 

A recent floristic survey of Edmonson and Warren Counties has 
revealed the presence of two species of this genus. Isoetes engel- 
mannii has been collected from three different localities in Edmon- 
son County, and 7. melanopoda Gay & Durieu has been collected 
from wet prairies and temporary pools in Warren County south 
of Bowling Green. 

Dr. Warren H. Wagner, Jr, has kindly examined the Warren 
County specimens and has stated that they definitely belong to 
the L melanopoda group. Voucher specimens have been deposited 
in the herbarium of Western Kentucky University, 

It is anticipated that as more extensive field investigations are 




Elmore, Department of General Biology, Vanderbilt University, 
Nashville. Tenn. 37203. 

PsiLOTTTM NUDUM New TO ARIZONA. — lu a collectiou of plants 
received for identification at the University of Arizona was a 
specimen of Psilotum nudum (L.) Palisot, a new record of the 
genus for Arizona. The specimen was collected by IXIr. Jack 
Kaiser, of the Plant Quarantine Service, U. S. Department of 
Agriculture, in Sycamore Canyon, Santa Cruz County, Arizona 
{Fig. 1), This canyon has been miich studied by Mr. Leslie Good- 
ding, who, however, never found this species. The plant grows in 
rock crevices {Fig, 2) and is definitely not plentiful. I have made 
several collecting trips to search for it, and I did find the one 
location where it grows. However, since it is so rare, I collected 
only just a small sprig or two for the University of Arizona Her- 
barium. The canyon is steep and hard to negotiate, but a careful 
search has not turned up any additional plants. Sycamore Canyon 

American Fern Journal 

Volume 60, Plate 8 



32 American Fern Journal 

is located about 20 miles west of Nogales. The location for the 
Psilotum is about a six mile hike down the canyon. 

Psilotum nudum is widespread in the tropics and occurs also in 
Florida and the southeastern United States. It has been recorded 
also from Texas by Correll, and from the Rio Mayo, Sonora,. 
Mexico, by Gentry. The latter is the closest recorded locality. 
Incidentally, the Ball Moss, Tillandsia recurvata L., the only 
epiphytic flowering plant known in Arizona, also grows in Syca- 
more Canyon and a few other southeastern Arizona localities. It 
has a range similar to that of Psilotum — Florida and the south- 
eastern United States, Texas, and northern Mexico.— Walter 

S. Phillips, Department of Biological Sciences^ University of 
Arizona. Tucson. Arizona R5721 . 

Notes and News 

The American Fern Society Annual Meeting this year will 
be with the A.I.B.S. at Bloomington, Indiana, on the campus of 
Indiana University. Dr. Charles Heiser will be our local representa- 
tive. We will have a foray on August 20-22. Foray reservation 
requests should be sent to Miss Lois Carlson, Matthaei Botanical 
Garden, University of Michigan, Ann Arbor, Mich, 48105. On. 
August 24 we will have a Society luncheon, along with a program 
of papers. Dr. John T. :Mickel, New York Botanical Garden,. 
Bronx Park, Bronx, N. Y. 10458 is the program chairman. On 
August 25 the Society will sponsor a symposium on 'Terns as 
Tools in Solving Biological Problems," The General Section and 
the Developmental Section of the Botanical Society of America will 
co-sponsor the symposium. 

Recent Fern Literature 

The Illustrated Flora of Illinols: Ferns, by Robert H. 
Mohlenbrock, Southern Illinois University Press, Carbondale^ 
Illinois, 1967. 119 pp. $8,00.— This, the most recent state fern 
flora, is similar in format to Billington's ^Terns of Michigan,'' but 

Recent Fern Literature 33 

it does include some new data regarding spores and chromosome 
numbers not treated by Billington. It differs from VannorsduU's 
''Ferns of Ohio" in that it is illustrated by drawings rather than 
photographs, which is not always an advantage. Aluhlonbrock's 
drawings of details are good, as are some of those of small ferns, but 
the larger compound ferns like Dryopleris carthusiana (Z). spinu- 
losa), D. intermedia, and others are not recognizable from these 
drawings but are readily recognizable from Vannorsdall's photo- 
graphs. The last enumeration of Illinois pteridophytes recorded 67 
species; the present work treats 81 species and six varieties. 
This large increase is partly due to intensive fieldwork during the 
last few 3^ears and partly due to different specific concepts. The 81 
species include a number of hybrids perhaps not qualifying as 
true species. The treatment of each species includes the common 
names, habitat, general range, Illinois range with map, and taxo- 
nomic comments. One new variety is described, Lycopodium 
lucidulum var. tryonii, which differs in having the leaves entire, 
and which is said to be known only from Jackson County, Illinois. 
It seems rather unlikely that a true variety of the widespread L. 
lucidulum would be known from a single locality. The plant de- 
scribed could well be a hj'brid with L. porophilum, which appar- 
ently has not been collected in Jackson County, but which does 
grow not far away. I believe that I have heard of reports of such 
entire-leaved plants from outside Illinois also. There are a few 
errors that might be pointed out. The drawing of Adiantum 
pedatum is inaccurate in not showing the solitary pinnules that 
are present on the internodes beyond the second fork; these 
isolated pinnules are important for they maj' help to explain the 
branching, if they are equivalent to all the rest of a branch. 
Dryoptens cristata is said to have a chromosome number of 2n = 
246, but I believe that this is an error, for this is regarded as a 
tetraploid (2n = 164} rather than a hexaploid. The description of 
the indusium of Dennstaedtia as '^modified from a recurv^ed tooth of 
the leaf margin" is not accurate, for the indusium is partly an 
extrorse inner indusium united with a modified introrse marginal 
lobe. The entry in the key on page 85 separating Woodsia from 

3-i American Fern Journal 

o W. ilvi 


these are strictly minor criticisms. The book is creditabJe and 
usable and should be in the libraries of fern students.— C.V.M. 

Vascular Plants of the Pacific Northwest, Part 1. By 
C. L. Hitchcock, A. Cronquist, M. Ownbey, and J. W. Thompson. 
914 pp. 1969. University of Washington Press, Seattle, Washington 
98105. S25.00. This, the fifth and last volume of the flora of the 
Pacific Northwest, completes this monumental work, the first 
parts of which were pubhshed several years ago. Although the 
last to be issued, this is the first part, which contains the treatment 
of the vascular cryptogams, the gymnosperms, and the mono- 
cotyledons, including the large and difficult groups Cyperaceae 
and Gramineae. The present part contains a glossary, a general 
mdex, and an appended Hst of additions and corrections to the 
previously published parts. The ferns and fern aUies are treated 
by Dr. Cronquist, and very well too. The keys are accurate and 
usable not too technical but technical enough when necessary. 
One detail is that finite verbs are sometimes used in the keys 
e.g. p. 57, ". . . indusium which is so close to the margin . . ." or 
''submarginal row that is protected by . . ."; the use of such 
"which" clauses is not usual in botanical keys and is never neces- 
sary. The treatment is conservative in its recognition of varieties, 
species, and families. I approve of placing the true ferns all in the 
inclusive family Polypodiaceae at least until much more is known 
about their morphology, cytology, and evolution. Cronquist 
does indicate in the key some of the groups that have been called 
families, including one called "Aspidiaceae." This is an incorrect 
name because according to the Code of Nomenclature a family 
name must be based on the name of a legitimate genus, and Aspi- 
dium is an illegitimate generic name, superfluous when published 
because it originally contained the type species of several earlier 
generic names that ought to have been adoped, such as Dnjopteris, 
Thelypteris, Tectaria, Athyrium, Polystichum, and perhaps others. 

Recent Fern Literature 35 

The proper family for this group depends on how finely it is split 
up itself into families; in the rather inclusive sense used by Cron- 
quistj the proper name is probably Aspleniaceaej but if Asplenium 
is placed in a different family, the remainder should probably be 
called Dryopteridaceae. The names of the classes accepted by 
Cronquist are Lycopodiatae, Isoetatae, Equisetatae, and Poly- 
podiatae, according to a suggestion that Cronquist, Jeffrey, and 
Takhtajan made to the recent International Botanical Congress. 
However, the Congress rejected this change, and so these names 
should not be accepted, since the Code says that the termination 
for classes should be '^-opsida." The '^ Polyp odiatae" should prob- 
ably be called the Filicopsida. SeJagineJla is included in the Isoe- 
tatae^ presumably chiefly because of its heterospory, but I can't 
really beUeve that Selaginella is more closely allied in a phylogenetic 
way to Isoetes than to Lycopodium, with which it seems obviously 
to have much more in common, Cronquist includes the hetero- 
sporous Azolla and Marsilea in the Polypodiatae, and so there is 
no a priori reason why heterospory could not have developed 
also within the Lycopodiatae. Types are cited, as they should be 
in a flora when space permits, but they are done here in a verj^ 
unusual way. Normally a type is cited either after the accepted 
name or after the name-bringing basionym, and so it is here 
sometimes. But not always. For instance, in Polypodium hesperium 
Maxon (1900), the type is not cited after this original and accepted 
name but following the later synonym P. vulgare var, hesperium 
Xels. & Macbr. (1916); there does not seem to be any justification 
for this. 

There are a number of details that ought to be commented on. 
The author of the combination Cystopteris montana (Lam.) is 
left undecided; the authority ''Bernh." used by Blasdell in his 
revision and by others is wrong; however, there seems to be no 
reason not to accept Desv, Mem. Soc. Linn. Paris 6: 264. 1827, as 
used in the 'Tlora Europaea," until someone can come up with an 
earlier usage, which is probably not possible. The authority for 
Blechnum spicant is given as Withering (1796), but it was shown 
long ago by :\Iansfeld (Repert. Sp. Nov. 45: 202. 193S) that the 

36 American Fern Journal 

combination had been made two years previously by Roth (in 
Usterij Neue Annalen 2(10): 46. 1794) (cf. Maxon, Amer. Fern J. 
34: 51. 1944). The authority for Woodicardia fimhriaia has been 
accidentally omitted; it is J. E. Smith in Rees; incidentally, I 
saw and photographed the holotype in the J. E. Smith Herbarium 
in the Linnean Society last spring. It is the stated intention to 
include all synonyms based on material from the Pacific North- 
west, but one has been omitted here — Blechiium doodioides Hook. 
Fl. Bor. Amer. 2: 263. 1840. The type locality of this species is 
not definitely known, but it was collected by Douglas and surely 
in the Pacific Northwest; Hooker thought that it might have come 
from Hudson Bay, but that is obviously impossible since this 
species does not grow there or anywhere nearby. This synonym is 
of some importance, for it would be the earUest specific epithet for 
the plant of this area if it should ever be separated from the 
European type, which is not at all inconceivable. The type was a 
somewhat abnormal specimen with the sori somewhat interrupted- 

Another synonym that has been omitted, although the type is 
from the area concerned, is Woodwardia paradoxa C. H. Wright 
(Gard. Chron, 41: 98. 1907), a synonym of W. fimhriata. The 
type w^as collected on a small, undesignated island off the coast of 
Vancouver Island, B. C, by F. W. Moore, and wall presumably 
be in the herbarium at Kew\ Strangely enough, Wright did not 
compare his species with W. fimbriata or its synonym W. chamissoi, 
but with the European W. radicaiis, apparently unaware that the 
Pacific Coast plant had ever been separated as a species. 

Cheilanthes lanosa Michx. is reported for the northwest for the 
first time in this publication. The plant is supposed to have been 
collected at high elevations on INIount Jupiter in the Olympic 
Mountains of Washington. I have been aware of it for several 
years, but I have had some doubts about it. This is an eastern 
species and the nearest known natural occurrences are in 
Minnesota, more than 1,000 miles to the east. The plant on Mount 
Jupiter was brought down and cultivated in a garden in Brinnon, 
Washington, and it is possible that here it became confused 
with a cultivated plant of the eastern C. lanosa. Later attempts to 

Recent Fern Literature 37 

find this species again on Mount Jupiter have been unsuccessful. 
Following the '*Flora Europaea*' and some recent authors, Cron- 
quist has adopted Isoetes setacea Lam, for the common spiny- 
spored plant of the northern United States and Canada. This has 
very recently been shown to be wrong; the comment by C. Jermy 
(Brit. Fern Gaz. 10: 106. 1969) deserves to be quoted in full: 
^^Isoetes setacea Lam. for L. echinospora Dur. is an unfortunate 
change for which I am partly responsible. When I wrote the 
account of Isoetes for ''Flora Europaea," I was aware that Roth- 
maler and later Fuchs had identified the type specimen of /, 
setacea Lam. as /. echinospora, I had seen only a photograph and 
been told that the plants were immature, and I had agreed that, 
superficially, this specimen did indeed have the habit of /. echino- 
spora. HoAvever, I have since had occasion to study this specimen 
in the Paris Museum and through the kindness of Mme. Tardieu- 
Blot I was able to remove a few immature megaspores and study 
them under the scanning electron microscope. Only now, after 
seeing some hundreds of megaspores from a wide range of species 
and at various stages of maturity, do I feel confident in placing 
the Lamarck specimen in the section Tuberculatae and not Echina- 
tae. In other words we must revert to the status quo and I must 
apologize to ]\Ir. Harrison and others who have taken up the name 
J. setacea Lam., the identity of which is now^ being investigated." 
This illustrates the coming importance of the scanning micro- 
scope for spore studies. The proper name for the Pacific Coast 
plant called /. setacea by Cronquist is probably 7. echinospora Dur. 
Recently, Fuchs (Nova Hedw. Beih. 3: 51-78. 1962) has taken up 
the earlier name 7. tenella Desv. (^lem. Soc. Linn. Paris 6: 179. 
1827), but the identity of Desvaux' type (FL Danica L 191) with 
I. echinospora has been challenged. As a variety the name is 7. 
echinospora var. braimii and as a distinct species 7. muricata Dur. 
The species Dryopteris austriaca Jacq. is accepted in the inclusive 
sense that I myself adopted in my treatment of the eastern ferns 
in Gleason's '*New Illustrated Flora'' (1952). I still think that this 
is a possible treatment of the group, but recent cytological studies 
have indicated that it might be better to recognize several species. 

38 American Fern Journal 

If so, the position in the Pacific Northwest is far from clear. Most 
of the plants would fall into what has been called a diploid Dry- 
opteris dilatata. But the true dilatata from Europe (D. austriaca 
Jacq. var. austriaca) is a tetraploid, and it has been suggested that 
our Pacific coast dilatata is really the diploid species D. assimilis S. 
Walker (in Clapham, Tutin, & Warburg, Brit. Fl ed. 2. 1962) of 
the British Isles; this has been indicated not only by the chromo- 
some number but also by chromatography. The true tetraploid 
D. spinulosa does apparently also reach into the Pacific Northwest, 
for I have seen a specimen that seems to be surely this rather 
than D. assimilis (Orogrande Creek, Clearwater County, Idaho, 
Epling & Houck 9349). The proper name for "D. spinulosa*^ 
(an illegitimate name) is still uncertain; the name D. carthusiana 
(Vill.) H. P, Fuchshas been accepted recently, but Jermy indicates 
that there is grave doubt that Villar's type is really this species; 
this kind of doubt is to be expected in a group like this that is 
cytologically complex and little understood even after intensive 
study for many years. — C. V. M. 

A Revised Flora of Malaya, vol. II — Ferns of Malaya, ed, 
2, by R. E. Holttum, Gov't. Printing Office, Upper Serangoon 
Road, Sinagpore, 13, 1966. 653 pp. ca. $7.50 H-postage.— The orig- 
inal edition of this popular and useful book appeared in 1954 (see 
this Journal 46: 15S. 1956). Despite one reprinting, it has been 
out of print for several years. The new edition is largely identical 
to the first. A few pages, however, are totally redone and Appendix 
II, a ten-page hst of changes and corrections to the first edition, 
is added. In recent years several studies bearing on the Malayan 
fern flora have been published, principally by Prof. Holttum and 
Mrs, B. E. G. Molesworth Allen. Besides incidental changes in 
many genera, considerable emendations have been made \i\Cijathea, 
Dicranopteris^ Gleichenia, Lygodium^ and Ophioglossum, and 
Diplazium has been separated from Athyrium, Entries in the main 
text that are amplified or corrected in Appendix II have been 
marked with an asterisk. Names in Appendix II unfortunately 
have not been included in the index. For readers who have the 

Recent Fern Literature 39 

first edition, Appendix II alone may be purchased from the 
publisher for about $0.7o. — D.B.L. 

Chromosomes and Speciation in FEUNb, by Kunio Mitui. Sei. 
Rep. Tokyo Educ. Univ. [Tokyo Kyoiku Daigaku], B, 13 : 285-333. 
19GS. — This is the first comprehensive listing of chromosome 
numbers of the Japanese ferns, treating 66 genera and 202 species. 
The mean sizes of the chromosomes of some of the species are 
stated; these vary from 3 /i to 7,5 /x, A number of presumed 
triploids, autotetraploids, and allotetraploids were authenticated 
cytologically, some of which are distinguishable morphologically 
by spore size, stomatal size, texture, and habitat. Most homo- 
sporous ferns have relatively large chromosome numbers compared 
with phanerogams; Mitui believes that most modem diploid 
species are in reality old polyploids derived from ancient and 
extinct species with lower numbers. He considers that a similar 
chromosome size often indicates relationships among the genera, 
that the older genera have higher base numbers, that advanced 
genera have many polyploid or apogamous species, and that 
primitive genera like Dennstaedtia have many aneuploid species. 

C. V. M. 

BiBLioGR.\PHY TO Floras OF SOUTHEAST AsiA, by Clyde F. Reed. 
Published by the author, 10105 Harford Road, Baltimore, Md. 
21234. 1969. 191 pp. $3.50.— :M err ill and Walker's usefuP^A BibU^ 
ography of Eastern Asiatic Botany (1938, Supplement, 1960) 
covered the essentially temperate parts of eastern Asia from China 
and Taiwan northward, but not tropical southeastern Asia. This 
new index covers Burma, Laos, Thailand, Cambodia, Viet Nam, 
Malaya, and Singapore. The title is a httle misleading because the 
work includes not only floras, which are rather few, but chiefly 
smaller taxonomic papers and also papers on fossil, cultivated, 
agricultural, and medicinal plants, forest timbers, and taxo- 
nomically oriented ecological papers. The bibhography would 
be more useful if it contained a subject index. — C- V. M. 

40 American Fern Journal 

American Fern Society 

Report of the President for 196Q 

It is now time to give an accounting of our stewardship for the 
past year, but first I wish to extend my greethigs to all of our 


members, both old and new. To the new members w^e say- 
come involved in the society and take advantage of the various 
services available to you. Let us know your needs. 

In this report I shall only touch brieRy on some aspects of the 
Society because other council and committee members will have 
more details to add. 

As you all know^; the big event of the year w^as the fern field 
trip conducted prior to the regular meetings of the XI Inter- 
national Botanical Congress held in Seattle, Washington. This w^as 
a wonderful experience for those of us who had not previously 
botanized in the Pacific Northewst. It was a tremendous thrill 
for me to see Cheilanthes gracillima and Aspidotis densa (formerly 
Cheilanthes siUquosa) growing in all their glory, since these are part 



ship of the Fern Spore Exchange but also for arranging a wonderful 


Over 80 people attended, including some members from here and 
abroad whom we had not seen in some years. 

Our next meeting w^ill be in conjunction with the American 

Institute of Biological Sciences next fall in Indiana. We plan a 




America in Edmonton 

fern flora is richer in Colorado than it is in that part of Canada. 

Parts of the booklet on ferns have been written and we expect 
further progress this coming year. The Journal Index and member- 
ship list are also being worked on by committees set up by your 
President, May I again issue a plea to all of our members to 

American Fekx Society 41 

remember the Society financially in whatever way seems appropri- 
ate. We will need more money than we can now spare for the above 
publications. In addition, we would all like to see a larger number 
of pages in the Journal, and this costs money. Mrs. Una Weatherby, 
Miss Scamman, Dr, Wherry, and others have made gifts or be- 
quests in the past, and we hope their example will be followed. 

When one attempts to assess the health of the American Fern 
Society by checking membership figures, one finds an up-and-down 
pattern during the past ten years. We now have about GOO domestic 
and foreign members plus over 300 institutional subscribers to the 
Journal. In recent years the low was 633 members in 1968 and the 
high was 896 in 1962. When I consider the general increase in 
population in the United States, and when I view the above 
figures, I can only say that I am not too pleased with our member- 
ship growth. Your Council grapples with this problem every year. 
Do you have a suggestion for increasing membership? Would each 
of you try to get one new member this year? If you are organization- 
minded and live in a fern-rich area, why not contact Society 
members and friends and plan a local field trip? We certainly 


the health of the Society. 

submitted, Irving W. Knobloch 

Report of the Secretary for 1969 

Individual membership in the American Fern Society at the end 
of 1969 stands at 603 members, with an additional 327 institutional 
snbficribers to the Journal. ^Members and subscribers come from 

every state m 
figures repress 


896 in 1962. This may be a time to pusn 
evaluate our role and function as a society. 



icArthur, a member since 1964 
member since 1953. 

42 A:\iERicAN Fern Journal 

The annual meeting of the Society was held in conjunction with 
the XI International Botanical Congress, August 24-September 
2, 1969, at The University of Washington in Seattle. The Society 
joined with the Congress in sponsoring a Pre-Congress Pteridology 
field trip from August 18 to August 22 in the Cascade ^Mountains 
of Washington. This trip will be reported in detail at a later date. 
Because of the decision of the Congress that there would be no 
sessions devoted to any special plant groups, there was no formal 
session at which members of the society presented research papers. 
Rather, research papers concerning ferns were scattered through- 
out the program of the Congress; it is impossible to list them here. 
The formal meeting of the Society was a luncheon meeting on 
Tuesday, August 26, attended by about 80 persons and presided 
over by President Irving Knobloch. Several foreign members 
attending the meeting were Dr. H. P. Fuchs from the Netherlands, 
Dr. R. E. Holttum from England, E. A. Turnau from Malaya, and 
several from Canada, including Dr. T. M. C. Taylor and Airs, E. 



with all members 

present. Approval was given to reprint copies of 20 back issues of 


funds from the Greenpoint Savings Bank to an appropriate bank 
in Pittsburgh which would afford the Society a greater interest rate. 
Discussion also concerned progress on a booklet on ferns for the 
general public. It is hoped that this booklet will soon become a 
reality, as it would be a great help to the Secretary in responding 
to numerous requests for general information on ferns. 

Respectfully submitted, A. Murray Evans, Secretary 

Report of the Treasurer for 1969 

Although receipts from all sources for the year 1969 were 
$7,136.83, which is much lower than last year, we still ended the 
year with $2,617.68 in the checking account. We invested the 
Scamman bequest of $3,000 in a savings account, where it will 
draw 5 per cent interest compounded quarterly. Our four savings 

American Fern Society 



S85.22, and sale of reprints by $4C) 


of the Journal from the checking account. 




or resignation, making a gain of 7. 

Taking into consideration the above statements, I believe that 
the American Fern Society is in good financial condition. 


Cash on Hand, January, 1969 
^Membership Dues 








Advances — 1970 
Advances — beyond 1970 


Sale of Back Issues of Journal 
Sale of reprints 

Royalties (Wherry's books) 

Page and Plate charges 
Advertising in Journal 
Spore Exchange for 1968 


Total Receipts 









S 6,902.15 














44 American Fern Journal 


American Fern Journal 

Vol. 58, No. 4 1,032.20 

Vol. 59, No. 1 960.35 

Vol. 59, No. 2 1,058.93 

Vol. 59, No. 3 1,198.96 


Reprints and postage 445.37 

Envelopes for mailing Journal 97.00 

Printing stationery, envelopes, billheads, etc. 100.25 

Treasurer's expenses (stamps and supplies) 111 . 14 
Secretary's expenses 
Editor's expenses 


Ad of Society in L. A. Fern Publ. 10.00 

Treasurer's bond renewal for 3 years 31 .00 

Deposited in Green Point Sav. Bank (Life Mem. Fund) 80.00 
Opened a Savings Account at Union Nat. 

Bank (Scamman Fund) 3,000.00 

Refunded to Agencies 
AIBS dues 

400 . 00 

Reprinting of back issues of Journal 2,860.00 



Cash on hand, January 1, 1970 $ 2,617.68 

Statement, December SI , 1969 


Cash in Union National Bank $ 2,617.68 

Cash in Green Point Savings Bank 


Bissell Herbarium Fund 
Life Membership Fund 
Weatherby Fund 

Cash in Union Nat. Bank 
Scamman Fund 

Accounts Receivable 


(Two sets of Journals and 6 reprint bills) 600.54 

Inventory, Journal (23,178 @ .50) 11,589.00 

Library' 396 . OO 



American Fern Society 45 


Advance dues 

Advance Subscriptions 

Fund balances 

Bissell Herbarium Fund 
Life ilembership Fund 
Una Weatherby Fund 
Scamman Fund 

General Fund 








Respectfully submitted^ 

LeRoy K. Henry, Treasurer 


Report of the Auditing Committee 

We hereby certify that we have seen the books and accounts of 
Dr. LeRoy K. Henry, Treasurer of the American Fern Society, Inc., 
and have obtained confirmation of the correctness of the Society's 
balance on hand as set forth in detail in the accompanying report 
of the Treasurer. 

Dorothy L. Pearth, Auditor 

Report of the Judge of Elections 

The essential results of balloting for officers of the American 
Fern Society are as follows: 

For President: 

Warren H. Wagner, Jr. 280 

For Vice-President: 

John T. Mickel 278 

For Treasurer: 

LeRoy K, Henry 276 

I therefore declare the following candidates elected to office: 
Warren H. Wagner, Jr., President; John T. ]Mickel, Vice-President; 
LeRoy K. Henry, Treasurer. 

Respectfully submitted, Alan R. Smith, Judge of Elections 

46 American Fern Journal 

Report of the Fern Spore Exchange 

The Fern Spore Exchange has made consistent growth in all 
respects. Twenty three universities or botanical gardens of other 
countries are participating in the Exchange, along with 65 in the 
United States. The fern societies of England and Japan both 
contribute to the Exchange, as well as many individuals through- 
out the world. 

A supplemental list to the 1967 and 1968 list has been distributed. 
These two lists include 489 species and varieties of ferns. Many 
of the items in the 1967 1968 list have been replaced with fresh 


450 to 939. 

The packets of spores distributed in 1969 were almost double 
the number sent out in 1968, but the packets received were about 
the same as in 1968. The Berlin Botanic Garden and the University 
of California at Berkeley contributed a total of over 130 rare, 
choice species. 


L. Wigj 

arranged for duplicating the lists over the past several years. 

Please collect and send in fresh spores or fertile fronds. The 
Exchange must provide /res/i spores at all times. Some items on the 
original list were collected in 1967 and some are out of stock. 
Therefore, the exchange is grateful for spores of all species. 

Respectfully submitted, Neill D. Hall, Director 

New Members 

M. Acedo-Diaz, Ricarda-Hiich Strasse 6, 5 Koeln-80, Germany 

Mrs. Virginia Ault, 1030 Oriole Ave., Miami Springs, FL 33166 

Mr. John J. Barnes, 26 Bertmore Dr., Stamford, CT 06905 

Mrs. Samuel M. Beattie, 30 Woodland Way, Greenville, SC 29601 

Mr. Eugene A. Bond, West Road, Dorset, VT 05251 

Mrs. P. N. Booker, Jr., 103 Pine Tree Dr., Gulf Breeze, FL 32561 

2vlr. Woodward S. Bousquet, 44 Waverly St., Pittsfield, MA 01201 

Mrs. Laurel A. Cawrse, 21793 Mastic Rd., Fairview Park, OH 44126 

Mr. James R. Falender, 11-5 Edgehill Terr., Troy, NY 12180 

American Fern Society 47 

Mr. Scott Fikes, 3220 Botanic Garden Dr., Ft. Worth, TX 86107 
Mrs. Philip D. Fletcher, Unicoi, TN 37692 

Mrs. Richard O. Grisdale, 33 W. Beechcroft Rd., Short Hills, NJ 07078 
Mr. Goeffrey R. Ilalliwell, No. 9 The Avenue, Bellanibi, 2518, N.S.W , 


Mr. William Harper, Lena High School, Lena, WI 54139 

Mr. J. M. Jacobsen, SSO E. Snell Rd., Rochester, MN 48063 

Mr. Leopold W. Kaiis, 1634 E. 96th St., Brooklyn, NY 12336 

Mr. Jack Langrell, 9257 29th St. W., Seattle, WA 98126 

Mrs. Madeline Lovett, 6016 Westchester Dr., College Park, MD 20740 

Mrs. Harry D. Lubrecht, 4672 Broadway, New York, NY 10040 

Mr. Robert Magill, 208 Sul Ross Village Apt., Alpine, TX 78930 

Mr. Joseph F. Major, 47 Highland Ave., Somervllle, NJ 0S876 

Mr. ]\Ionte Manuel, 9610 Lawndale Ave., Tacoma, WA 98498 

Mr. John E. McNeal, 5757 Country Club Dr., Oakland, CA 94618 

Mrs. Edith Meyer, Rt. 3, Box 658, Fort Myers, FL 33901 

Miss Judith G. Morgan, Dept. of Botany, Univ. of Texas, Austin, TX 78712 

Dr. Takehisa Nakamura, Botanical Laboratory, Tokyo Nogyo Univ., 1-1, 

Sakuraga Oka, 1-chome, Setagaya, Tokyo 156, Japan 
Mr. Isadore Nicholson, C. W. Post College, Long Island University, Green 

vale, NY 11548 
Mr. Robert E. Oman, 3011 N.W. 18th Terr., Miami, FL 33125 
Major C. Rhodes, Spartan High School, Biology Dept., Spartanburg, SC 29302 
Rev. Austin A. Rodgers, 2624 S. Lincoln Ave., Springfield, IL 62704 
Mr. Richard I. Sindell, 11271 Champagne Point Rd., Seattle, WA 9S033 
Mrs. Calvin Skinner, Dorset, VT 05251 

Mrs. Bruce W. Spaulding, 126 Cedarwood Rd., Stamford, CT 06903 
Mr. Wayne M. Taylor, R.R. 2, Olney, IL 62450 
:Mrs. Marcelle Thompson, Rt. 4, Box 154, Whitesville, NC 28472 
Miss Marguerite A. Weaver, Box 134, Glyndon, MD 21071 
Mrs. Bell L. West, 2744 McDowell St., Durham, NC 27705 
Mrs. \'erna H. Wilkinson, 2732 Haverford Place, Charlotte, NC 28209 


Changes of Address 

IS. Elva R. Beard, P.O. Box 862, Salem, VA 24153 

Dr. Dorothy C. Bliss, Randolph-Macon Woman's College, Box 278, Lynch- 
burg, ^'A 24504 

Mrs. Caroline E. Carter, Dept. of Anthropology, Duke Univ., Durham, N.C. 


Mr. Richard B. Chillas, Jr., 7141 McCallum St., Philadelphia, PA 19119 

Aliss Carol E. Copeland, Biology Dept., Univ. of Tennessee, Chattanooga, 

TN 374 03 

48 American Fern Journal 

Mrs. Blanche E. Dean, P.O. Box 14, Goodwater, AL 35072 

Mr. Arthur Decker, P.O. Box 10 04 42, 5 Koehi-1, Germany 

Mr. Kenneth J. De Nault, 125 Miramontes Rd., Woodside, CA 94062 

Mrs. Marie EUcker, 2224 Shelby St., New Albany, IN 47150 

Mr. Harold W. Elmore, Dept. of Genl. Biology, Vanderbilt Univ., Nashville, 

TN 37203 
Mrs. Lorraine Ennemoser, 14221 S.W. 86th Ave., Miami, FL 33158 
Dr. Donald R. Farrar, Dept. of Botany & Plant Path., Iowa State Univ., 

Ames, lA 50010 
Mrs. George N. Gardner, Piper Hill Rd., Weston, VT 05161 
Mrs. John Zell Gaston, Box 36798, Sharpstown Station, Houston, TX 77036 
Mr. Jack W. Gilbert, 8607 Timberside Dr., Houston, TX 77025 
Miss Nora Sue Hollis, 3709 Walnut St., Kansas City, MO 64111 
Mr, Victor J. Larson, 8431 W. North Ave., Wauwatosa, WI 53226 
Mr. J. Harry Lehr, 12434 N. 25th Place, Phoenix, AZ 85028 
Dr. Jewel Moore, Box 915, State College of Arkansas, Conway, AR 72032 
Mr. Hideaki Ohba, No. 4, Kitasuna 5-10, Koto-ku, Tokyo 136, Japan 
Miss Aleta Jo Petrik, Biology Dept., John J. Pershing College, Beatrice, NB 

Maj. Peter G. Root, 431 St., Medical Co., Ft. Sam Houston, TX 7S234 
Miss Hester M. Rusk, 40 Riverside Ave., Red Bank, NJ 07701 
Dr. Arthur L. Shuck, 207 N. 6th St., Elsberry, MO 63343 
Mr. Franklin D. Snyder, P.O. Box 901, Toccoa, GA 30577 
Dr. Raymond E. Stotler, Botany Dept., Southern 111. University, Carbondale, 

IL 62901 

Exotk and Hardi Ferns 



2131 Volleio Street 
St. Heleno, Colifornio 94574 

Open Saturdavt and Sundays from 10 A.M. to 4 P.M. and bv aDDoinfment 

963-2998— Area 


Ownership, Management and 


Office, as established under the act of OctoI:>er 23, 1962, Section 4369, Title 
39, United States Code, the following statements are published. 

Title: American Fern Journal 

Frequency of Issue: Quarterly (Approximately March 31, June 30, 

September 30, and December 31) 

Location of Office of Publication (Printers): 3110 Elm Avenue 

Baltimore, Maryland 21211 


K. Henry, Section of Plants, Carnegie Museum, Pittsburg, 
Pa. lo213 


Museum, Pittsburg, Pa. 15213 

Editor: Dr. David B, Lellinger, Department of Botany, Smith 



American Fern Society, Inc., Section 
Museum, Pittsburgh, Pa. 15213 


Total number of copies printed: 
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The statements made above are certified to be correct, and are signed by: 
David B. Lellinger, editor-in-chief of the American Fern Journal on October 
1. 1969. 


VOL. 60 April- June, 1970 No. 2 

l^merican Jfern Sfournal 


Publifthad hy thm 









The Fern Collections in Some European Herbaria, VI. . . .C. V. Morton 49 

Fertile Sporophytes of Botrychium Tirginianum Attached to Gameto* 

phytes Bruce W, McAlpin 62 

A PecuGar Species of Grammitis C.V.Morton 65 

Vascular.Organization of the Rhizome of Cibotimn barometz 

Subhash Chandra 68 

A Surrey of Some Morphological Features of the Genus Elaphoglosstmi 

in Costa Rica Robert M. Lloyd 73 

Notes and News. * 83 

Recent Fern Literature - 84 

American Fern Society 


Mi99oy«> B-rrANicAC 

JUL 1 7 19^0 


®l)e American Jfern ^ocietp 

Council for 1970 

Warren H. Wagner, Jr., Department of Botany, University of Michigan, 
Ann Arbor, Michigan 48104. President 

JoHN^ T. MiCKEL, New York Botanical Garden, Bronx Park, Bronx, New 
York. 10458. Vice-President 

A. Murray Evans, Department of Botany, University of Tennessee, Knox- 
ville, Tenn. 37916. Secretary 

LeRot K. Henry, Division of Plants, Carnegie Museum, Pittsburgh, Penn- 
sylvania 15213. Treasurer 

David B, Lelunger, Smithsonian Institution, Washington, D, C- 1M560. 


J^ational ^otittf B&eptesientattbefS 

Warren H. Wagner, Jr., University of Michigan AAA.S. Representative 
RoLLA M. Tryon, Jr., Harvard University AJ.B.S. Representative 

Smetican Jf ern 3fwxnal 


Davio B. Lelli.\ger .Smithsonian Institution, Washington, D, C. 20560. 

C, V. Morton. ........ . .Smithsonian Institution, Washington, D. C. 20560. 

RoLLA M. Tryon, Jr. 

Gray Herbarium, Harvard University, Cambridge, Mass. 02138. 

Ira L. Wiggins Dudley Herbarium, Stanford University, Stanford, Calif, 


An illustrated quarterly devoted to the general study of ferns, owned by 
the American Fern Society, and published at 3110 Ebn Ave., Baltimore, 
Md. 21211. Second-class postage paid at Baltimore. The pages of the Jour- 
nal are open to members who wish to arrange exchanges; membership lists 
assist those interested in obtaining specimens from different localities. 

Matter for publication should be addressed to the Editor-in-Chief* 

Changes of address, applications for membership, subscriptions, orders 
for back numbers should be addressed to the Treasurer. 

Subscriptions $5.50 gross, $5.00 net (agency fee $0,50); sent free to mem- 
bers of the American Fern Society (annual dues, $5.00; sustaining membership, 
$10.00; life membership, $100.00). Extracted reprints, if ordered in advance, 
will be furnished authors at cost, plus postage. 

Back volumes $5.00 to $6.25 each; single back numbers of 64 pages or 
less, $1.25: 65-80 pages, $2.00 each; over 80 pages, $2.50 each; Cumulative 
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Etbrarp anli H^erbarium 

Dr. W. H. Wagner, Jr., Department of Botany, University of Michigan, 
Ann Arbor, Michigan 48104, is Librarian and Curator. Members may borrow 
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^pote €xc|)ange 

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£S Director. Spores exchanged and collection lists sent on request. 

^itti ants ^tqntsiti 

back issues of the Journal, and cash or other gifts are 
are tax-deductible. Inouiries should be addressed to th 

gfmerican Jfern Journal 

Vol. 60 Aimul-June, 1970 No. 2 

The Fern Collections in Some European Herbaria, VI 



The herbarium and lx)tanical garden of the Staatsuistitut fur 
Allgemeine Botanik in Hamburg are not well known or often 
visited by taxonomists, in spite of the herbarium being one of 
the larger of the world herbaria and one of great importance 
for certain areas. There were private gardens in Hamburg from 
early times. The Hamburg Academic Gymnasium was founded in 
1C13, and many of its early professors were interested in botany, 
especially Joachim Jungius (ir>S7-1657), who was appointed 
rector in 1G29. Sachs, in his history of botany, states that Jungius 
was the first in Germany, as Caesalpimis was in Italy, to combine 
pliilosophical thoughts on botany with the true observation of 
living plants. It seems clear that Linnaeus studied the works of 
Jungius in his youth. Linnaeus visited Hamburg in his travels 
and commented on the richness of the gardens there, which in- 
cluded even then 50 species of the South African genus Mesemhry- 
anthemnm and 45 species of Aloe. 

The credit for the first large private garden must go to J. X- 
Buek, who was referred to as "\he German Philip Miller/' in 
reference to the noted EngUsh gardener of the same period. 
Buek's catalogue of 1779 htid 200 pages of plants for sale. 

Johannes Fluegge (1775-lSlC) was a talented Hamburg botanist. 
He got his doctorate in Erlangen in ISOO under Schreber, but even 
prior to this he had made many collecting tri{)s in Germany with 

Volume i\{}, No. 1, of the JorRN.VL, pp. 1-4S, was is>ued April 21, 1970. 

50 American Fern Journal 

his friend Floercke (himself later professor of botany in Rostoek 
and a noted bryologist of his time). He took the opportunity of 
making friends with all the botanists of his day — Martius, von 
?^roll, Jacquin, Schrader, Hedwig, Schkuhr. T.ater he travelled 
in France and Italy, collecting, studying, and making the acquaint- 
ance of Richard, Gouan, Gilibert, Jussieu, Desfontaines, Allioni, 
Bellardi, and others. He thought to work on a flora of France, but 
abandoned the idea when he found DeCandolle engaged on the 
same project. During these years he exchanged thousands of 
plants with botanists in Germany, France, and England. 

Fluegge returned to Hamburg in 1810 and decided to start a 
true public botanical garden. He sold about a hundred shares at 
400 Marks Banco (000 RM) and was able to buy Buck's establish- 
ment, which at the time included about 1200 species. Willdenow 
sent plants from Berlin, Sprengel from Halle, Hornemann from 
Copenhagen, Thouin from Paris, Fischer from Gorenki, Balbis 
from Turin, and Bernhardi from Erfurt, and soon there were 
3000 species in the garden, including many rare plants from 
Africa and Australia. Unfortunately, the army of Napoleon 
passed through Hamburg in 1813, leveling much of the city, 
including Fluegge's garden, which was completely destroyed, 
including the herbarium and manuscripts. Fluegge never recovered 
from this shock and died shortly thereafter of a ''nervous fever.'' 
He is remembered today only from his ''Graminum ■\Iono- 
graphiae'*, only one part of which was published (mostly on 
Paspalum), and the duplicate plants that he exchanged. 

Hamburg citizens, having had a botanical garden for a few 
years, were not content \^athout one, and so another was author^ 
ized. Johann Georg Christian Lehmann had been appointed 
Professor of Botany in ISIS, a post that had been vacant since 
Prof. Reimarus had fled Hamburg during the siege. Lehmann 
brought with him from Goettingen about 2000 species of plants 
for his private garden. When a site for the new public botanical 
garden was found near the Dammthor, these plants wore moved to 
the new garden. Lehmann himself planted the first tree XovembtT 
6, 1821. He brought the S^^'iss Ohlendorff from Bern as chief 

Ferns in European Herbaria ol 

gardener. The quaint, ^traw-thatched cottage erected for Ohleii- 
dorff was still standing and in use in 1900. 

Lehniann started the garden with a flurry of activity. Only two 
years later, in 1823, he sent out seeds of 52G4 species and received 
3609 packets of seeds and 2892 living i)lants. A coldhouse was 
built by public subscription in 1822. Lehmaiui continued his 
arduous duties for almost 40 years, not only as head of the garden 
but as the only professor of botany, and in addition at first he 
was obliged to teach zoology also. The number of species in the 
garden increased to 18,000 by 1848, and included many rare 
plants such as Dnjandra, Epacris, Borronia, WaUichia^ Xepeufhes, 
and Dionaea, Lehmann found time to continue his researches on 
his favorite genus Potentilla^ and also to publish ten parts of his 
'Tugilli,'' a series of papers describing interesting and new plants 
of the garden and herbarium. The later numbers of this were 
devoted almost wholly to Hepaticae. 

After Lehmann's death in 1860, Heinrich Gustav Reichenbach 
(1824-1889) was appointed Director. He was the son of the 
famous botanist Heinrich Gottlieb Reichenbach, Professor of 
Botany in Dresden. The son was interested in orchids from the 
first, and contributed the account of the Orchidaceae to his father's 
'Tcones Florae Germanicae et Helvetiae," on which he worked 
for ten years, with publication in 1851. He received his doctorate 
the next year by his thesis ''De pollinis Orchidearum genesis ac 
structura," which brought about his appointment as lecturer 
in the University of Leipzig, where he was made Extraordinary 
Professor three years later. He won the position in Hamburg over 

five competitors. 

On coming to Hamburg he made some fundamental changes, 
suspending the sale of plants to the public, eliminating the tree 
nursery and kitchen gardens, and starting a botanical museum. 
The plants in the garden had been arranged by the Linnaean 
system but he rearranged them by that of EndUcher, a much 
more meaningful arrangement. He began building up the collec- 
tion again, purchasing the entire collection of the famous English 
gardener Saunders, of Reigate, Surrey- 

52 American Fern Journal 

Reiehenbach continued his orchid studios, becoming the 
famous — and indeed ahnost the only — orchidologist of his time 
after the death of Lindley. He pubUished literally hundreds of 
papers in widely scattered journals, which included hundreds of 
new species. His name is usually abbreviated to ''Rchb. f/' by 
orchidologists, but this abbreviation is not in accordance with the 
rules as set forth in the Code. In this connection he travelled 
widely to various herbaria, often remaining for weeks or months. 
He was especially fond of Kew and planned on settUng there after 
his retirement. He is reported by contemporaries to have been 
jealous of others working on orchids, which he considered a 
private preserve. In an obituary (Proc. Linn. Soc. I.ond., August 
1891) a colleague wrote: ''A careful working of it [ Reiehenbach 's 
herbarium] will be needed to clear up the many puzzles in his 
descriptions, which of late years had assumed an esoteric charac- 


ter. . . . The state of his herbarium is quite unknown, for during 
long years past no botanist has been permitted to have even a 
glimpse of the collection. ... At his last visit to this country 
[England] Reiehenbach was looking forward to his retirement . . . 
and the preparation of a projected Index Reichenbachianus, which 
was to give a much-needed guide to the widely scattered descrip 
tious of species and varieties, but this last, if ever begun, is un- 
finished. At the Ghent Exhibition of ISSS it was noticed he was 
looking unwell and had shrunk from his accustomed portliness. 
By his death a gap is made which no man is likely to fill." 

Reiehenbach was an outgoing person, and his conversation was 
enlivened by witticisms and sarcastic remarks that were amusing 
to those who were merely onlookers and were not exposed to the 
brunt of his attacks. His sense of humor even carried over into his 
publications. He reported gleefully one story in the "Gardener's 
Chronicle:" It seems that a German entomologist described a 
butterfly under the name Sesia hngijormis. Reiehenbach wrote 
him that the name ''longiformis" was improper (because "formis" 


long if i 

"long" would be impossible). The entomologist dutifully renamed 

Ferns in European Herbaria 53 



to have remarked that if Reichonbach's herbarium ever came into 
his control he would make a funeral pyre of it. 

Reichenbach's chief claim to notoriety was his will, which even 
obtained notices in the newspapers, ordinarily unconcerned about 
the disposition of herbaria. In the London Evening Standard of 
June 5, 1889, appeared the following: "Professor Reichenbach': 
will is an event and a most disastrous one. It is indeed a scandal. 
That renowcd herbarium which gave to Hamburg, in the eyes of 
botanists, somewhat of the sanctity that moslems attach to the 
city where the Kaaba stands, is not only diverted from the use 
and destination which savants all over the world expected, more 
than that it is withdrawn from the service of mankind for twenty- 
five years. We do not know that Professor Reichenbach was a 
naturalized Etiglishman, but his labours and his fame were as- 
sociated with this country. It was taken for granted by men of 
science everywhere that the herbarium would go to Kew, and there 
is reason to think that he contemplated the bequest until very 
recent years. But neither Kew nor England is mentioned in the 
will. ... Of course there is a story connected with this most un- 
fortunate disposition— a story which is being discussed by all of 
the botanists of the world; but we would not dwell on it here. It is 
enough to say that the power of malicious individuals to damage 
the interests of the State is displayed in a striking manner." 

The will in question left his herbarium to Vienna on the con- 
dition that it remained unopened and sealed for 25 years from his 
death; if Vienna would not accept it under these conditions, it was 
to be offered in turn to Uppsala, the Gray Herbarium, and Paris. 

I do not know his reason for turning away from Kew. I asked 
Dr. John Hutchinson, who is the oldest working member of the Kew 
staff although of course not actively working in Reichenbach's 
time, but he did not know, other than to think that Reichenbach 
resented Rolfe having started to work on orchids at Kew. But I 
can guess at the reason for the peculiar provision in his will that 
the herbarium be sealed for 25 years. This coupled with the fact 

54 A^iERiCAN Fern Journal 

that he would allow no visitors to see his herbarium shows clearly 
that it was filled with fragments from various herbaria j those 
jokingly called *'kleptotypes/' By having the herbarium used it 
would be soon obvious that fragments came from Kew and other 
herbaria. After a lapse of 25 years most of the persons in charge of 
herbaria would be dead or retired and in no position to state 
whether the fragments had been removed by permission or not, 
That this guess is true is shown by the contents of the herbarium 
when it was opened, although perhaps no one has ever commented 
on the fact- The herbarium was found to be tremendous^ numbering 
more than 60,000 sheets of orchids and 8000 drawings. Almost 
all collectors of orchids are represented, including many from 
the late eighteenth centurj' and early nineteenth, such as Pavon, 

Commerson, Humboldt and Bonpland, Chamisso, Blume, Bory, 
Haenlce, Labillardiere, Martins, Reinwardt, and Gaudichaud.^ 
These specimens could hardly have come as duplicates to Reichen- 
baeh in such quantities and must have been removed from herbar- 
ium sheets, either with or without permission. This case is by no 
means unique. Christensen's fern herbarium purchased by the 
British Museum contains hundreds of fragments removed from 
various herbaria. The Underwood Herbarium, now in the New 
York Botanical Garden, contains hundreds of such ^'kleptotypes" 
also. The strangest case is that of Baillon, who worked at Paris 
for many years although not as a staff member. On his death his 
"type herbarium" went to Paris, where it is still maintained as a 
separate herbarium. It consists entirely of fragments removed 
from sheets in the Paris Herbarium, which is curious since the 
sheets in the Paris herbarium were freely available to him. These 
fragments really ought not to be considered the types of Baillon's 
new species but rather the full sheets from which they were re- 
moved. Baillon copied off the label data, and so the fragments 
can, at least usually, be matched up with the full sheets from 
which they were taken. 

*"\erzeichnis der im Orehideenherbare von Reichenharh fil. enthalteneTi 
Sammliingeii'' bv K. voii Kei."^sler and R. Rechiiiger. Aim. Naturhist. Hofmus. 
Wien 30: 13-23." 1916. 

Ferxs in European Herbaria 55 

Several years after Keicheiibach's death Dr. Zacharias (born 
1832 in Hamburg) became Director for many years. In 1901 all 
the botanical institutions were united to form the Botanische 
Staatsinstitut. In 1912 a separation was made into the Staats- 
institut flir Angewandte Botanik, with Dr. A. Voigt named 
Director, and the Staatsinstitut fiir Allgemeine Botanik, which 
included the Botanical Garden and the Herbarium Hamburgense. 
Dr. Hans Winkler became the first Director of the Staatsinstitut 
fiir Allgemeine Botanik. He was a student of Wilhelm Pfeffer, 
under whom he received his Ph.D. in 189S, working on physiology. 
He was instrumental in setting up the University of Hamburg in 
1919. His own research was on hybridization and parthenogenesis, 
and on apogamy and alternation of generations in ferns. He 
made collecting trips to the East Indies hi 1903-4 and 1924-5. 
His extensive Bornean collections were still partly unworked in 
1954 and probably still are; they have many undistributed dupli- 
cates also, I was told by Dr. Domke. Winkler retired during the 
War in 1943, when Hamburg was under constant attack. About 
sixty percent of the plants in the garden were destroyed by the 
bombing, but fortunately the herbarium escaped injury. Winkler 
died of a heart attack in 1945. In the same year Prof. Walter 
Mevius, a physiologist, was named Director. He was succeeded 
in 1964 by Prof. Horst Drawert, a cytologist and physiologist. 

Dr. Edgar Irmscher, a specialist on Begonia, was named Curator 
of the Herbarium Hamburgense in 1912; he served in that post 
until 1945. Dr. Walter Domke succeeded Irmscher. Since 19G1 
Dr. Kurt Walther has been Curator. Other staff membei^ of the 
herbarium of international reputation have been H. Hallier, who 
worked on Convolvulaceae, and C. Grimme, who worked chiefly on 
economic plants. 

The Herbabiu! 

In the earlv years of the botanical garden there was no official 

rM,+ .j11 tVip snecimens received 


in iiis personal herbarium, a practice common enough at one 
time but one to be deplored. Specimens that come to botanists 

56 American Fern Journal 

officially connected with an institution ought to be considered 
institutional property, and so they are today usually. But Leh- 
mann's herbarium was divided up into 150 different lots, which 
were individually sold and dispersed. We do not know today what 
it contained or where it went, and we are even more ignorant 
about it than about the herbaria of Kurt Sprengel or of Lambert, 
which were similarly broken up. It is perhaps not an easy matter to 
locate I.ehmann's types today. 

The beginning of the permanent herbarium was the gift of the 
private herbarium of H. W. Buek, of Hamburg, in 1864, which 
contained about 10,000 specimens. About the same time the algal 
herbarium of Nikolaus Binder was also donated. This is a large 
and important collection containing types or isotypes collected 
or described by Mertens, Tilesius, Agardh, Lyngbye, Raben, 
Hering, Montagne, Lindenberg, Rudolphi, Weber and :\Iohr, 
and many others. A little later the herbarium of the private 
Museum Godeffroy, built up by the Godeffroy family, was ac- 
quired. This contained the original collections of Graeffe, Amalie 
Dietrich, Kleinschniidt, and others who had been sent on expe- 
ditions by Godeffroy. The Graeffe collections are important for 
fern students, being the basis for Luerssen's "Filices Graeffeanae." 

The Buek and Binder herbaria were united in 1879, which is 
considered the official start of the present herbarium. Richard 
Sadebeck took over as custodium in 1882, and A. Voigt became 
his assistant. The herbarium grew rapidly until at the last esti- 
mation it contained 700,000 specimens. Sadebeck brought his 
own herbarium, mostly of European plants but containing also 
some exotics, such as Chilean specimens from Fhilippi. The 
grass herbarium of Klatt was acquired, and Willkomm's algal 
herbarium. But the largest accessions were of plants from Africa. 
The oldest of these and perhaps the most important were the 
South African collections of the Drege brothers Jean and Carl, 
who were born near Hamburg and who went to the Cape in 1826. 
They collected extensively, travelling by ox-cart to the mouth 
of the Orange River in the west and to Port Xatal (Durban) 
in the east, finding more than 7000 soecies. flbmit t.wn-+hirH« nf 

Ferns in European Herbaria 57 

all those that had been previously known in South Africa. The 
principal collections were in Berlin and were destroyed in World 
War II; there are only a few in Hamburg. Later Jean Drege was 
joined by Christian Friedrich Ecklon and Karl Zeyher, and the 
three of them continued to make extensive collections which 
were brought back to Hamburg to be studied in the nursery that 
the Drege family owned in Hamburg, Drege published a ^'Catalo- 
gus plantarum exsiccatarum Africac australioris" in Hamburg 
(1837-40), and Ecklon and Zeyher a more extensive scientific 
paper, "Enumeratio plantarum Africae australis extra tropicae," 
in three parts (1834-37). Many of the Ecklon and Zeyher plants 
^vere worked up by Otto Wilhelm Sonder. Sonder (born in Holstein 
in 1812) worked as a pharmacist for a living, but apparently all 
his spare time was spent on botany. He achieved an international 
reputation by his papers on Salix, on the algae of Australia col- 
lected by Preiss, and especially on the South African plants of 
Drege, Ecklon, and Zeyher, on which he worked with his friend 
Prof. William H. Harvey, of Dublin. The two were both interested 
m algae and proved congenial workers. Together they produced 
the fine three volumes of the ''Flora Capensis," published in 
1S59-1SC5. The work was left incomplete by the death of Harvey, 
but it is still a classic work on the South African flora. Sonder^s 
personal herbarium, the algae at least, was sold and is now partly 
in Stockholm and partly in the National Herbarium of Victoria, 

The more recent African collections constitute perhaps the most 
important part of the herbarium. They include the original 
set of Kurt Dinter's plants from Southwest Africa, the Max 
Julius Dinklage specimns from western Africa, especially Liberia 
and Gabon, the ^lildbraed collections from Cameroons (the 
3500 numbers of which are most important since the destruction 
of the first set in Berlin), plus collections by Fischer from Zanzibar, 
Stuhlmann from Mozambique (but not Tanganyika), Lindinger 
from the Canary Islands, and many others. A full account of the 
African collections has been published by Kurt Walther in his 
paper '^Afrikanische Pflanzen in Hamburg, Hamburger Botaniker 

58 A^iERiCAN Fern Journal 

ill Afrika" (Mitt. Geogr. GeselL Hamburg 56: 87-103. 19G5). The 
collections in the general herbarium are far too numerous to 
begin to enumerate them. 


The early works of the staff at Hamburg were partly published 
in the ^'Jahrbuch der Hamburgischen Wissenschaftlichen Anstal- 
ten," between 1889 and 1913, which was not however restricted 
to botany. The present serial publication is the ''Mitteilungen 
aus dem Institut fiir Allgemeine Botanik in Hamburg," volume 1 
of which appeared in 1914. The most important volumes in a 
taxonomic way are volume 7, which consists of the *'Beitrage 
der Flora von Borneo," a systematic treatment by families of the 
specimens collected by Winkler, with contributions by Diels, 
Kranzlin, Brotherus, Pfeffer, Pilger, Merrill, J. J. Smith, Chris- 
tensen (on the ferns), and Alston (on the Selaginellas). The intro- 
duction contains a map of Winkler's itinerary. Volume 10 (1939) 
is a ''Festschrift" for Hans Winkler on his 60th birthday; the 
principal paper is a "Flora des tropischen Arabien," by 0. 
Schwartz; I am not quite sure whether the material on which this 
paper is based is preserved in Hamburg or not, but it probably 
is, in which case Hamburg would be a necessary visit for anyone 
interested in the Arabian flora. This journal was interrupted by 
the war and did not resume publication until 1957. Only two 
volumes have been issued since then. 

The ^'Hamburger Garten und Blumenzeitung" was perhaps 
not quite an official publication, but it was closely associated with 
the botanical garden through its editor, Carl Friedrich Eduard 
Otto (1812-1855), usually called merely Eduard Otto. Otto was 
the son of the famous gardener of the Berlin Botanical Garden, 
Christian Friedrich Otto, and so he came by his gifts naturally. 
He started in as an assistant to his father in Berlin, but after the 
retirement of Ohlendorff he was chosen to come to Hamburg by 
Lehmann to be the head gardener, a post that he held for about 
40 years. The journal mentioned above was a gardening publi- 
cation, noticing plants new to cultivation and so forth, very 

Ferns in European Herbaria 59 

similar in content to the ''Gardener's Chronicar' concurrently 
beinoj published in England. Otto took over as editor with volume 4 
(184S) and continued through volume 40 (1884, the year before 
his death), altogether 37 volumes each with about 575 pages, with 
a total of over 21,000 pages. Since Otto did all the writing ap- 
parently, it is certain that he must have been industrious. The 
publication, like so many others, did not last long after Otto's 
death. This journal contains notes on many plants new to culti- 
vation and may contain some new names, although perhaps 
not intentionally with Latin descriptions. There are ferns dis- 
cussed, but I am sure that no one has ever looked through the 
volumes to see if any are new, nor is anyone likely to. The journal 
is by no means easy to read, being in cursive German type (except 
for the plant names in Roman type), and I have some doubt that 
many people ever actually read it even when it was being published. 
There is a portrait of Otto as a frontispiece to volume 41 of this 
journal, and a biography by F. Goeze (who took over as editor). 

Ferns in Hamburg 

The pride of the fern herbarium according to A. Voigt is the 

Prantl herbarium purchased after Frantl's death. Karl Prantl 
was born September 10, 1849, in Munich. He received his doctorate 
from Munich in 1870 under Xaegeli and Radlkofer. With his 
friend Lorentz, who later went to Argentina to teach and collect, 
he botanized in Bavaria and the Alps, and then spent a year as 
Xaegeli's assistant. In 1873 he was assistant to Sachs, with whom 
he worked on the regeneration of the growing point in angio- 
spermous roots. At Sachs' invitation he adapted some of Sachs' 
works into a "Lehrbuch der Botanik'' (1874), which became 
popular as a text in secondary schools and which went through 
many editions. In 1879 Prantl turned his attention to ferns, 
being at this time a professor in Aschaffenburg. His principal 
work in this field is his ^'Untersuchungen zur ]\Iorphologie der 
Gefasskryptogamen," I. ''Die Hymenophyllaceen" (1875) and 
IF *'Die Schizaeaceen" (1881). Prantl considered the Hymeno- 
phyllaeeae the lowest group of ferns from a developmental point 

60 A.MERiCAN Fern Journal 

of view. Subsequently, he worked on Cryptogramma, Pellaea^ and 
Ophioylossunij and his very last paper in 1892 was on fern classifi- 


cation. From 1889 until his death from tuberculosis in 1893 he 
was professor in Breslau. Adolf Engler evidently had a high regard 
for his talentSj for he asked Prantl to join him in a vast under- 
taking, a complete treatment of all the genera of plants, crypto- 
gams as well as vascular plants, something that had hardly been 
attempted since the time of Linnaeus. The work was to be called 
*'Die Natlirlichen Pflanzenfamilien," and Prantl was to be in charge 
of the cryptogamic portions, Prantl accepted willingly and began 
work at once, not on the cryptogams but on the early families of 
angiosperms according to the new system proposed by Engler. 
Prantl himself had no part in the elaboration of this system, 
which is now commonly called the Engler and Prantl system. The 
basic ideas of this arrangement of the families came from Eichler 
and were modified by Engler in his "Syllabus der Pflanzen- 
famiUen." Prantl worked up the Betulaceae, Fagaceae, ]Magno- 
liaceae, Anonaceae, Ranunculaceae, Berberidaceae, Cruciferae, 
and other families in short order, and it is apparent that had he 
lived he would have done a major part of the writing himself, 
something that Engler himself never did. Engler secured the 
cooperation of many botanists, chiefly German, to complete the 
work. Although Prantl died shortly after the start of the work, 
his name remained on the title page throughout. 

I have myself never had a high regard for PrantPs work on ferns. 
His idea of applying morphology and anatomy to taxonomic 
classification is certainly proper, but he seemed to lack the eye 
for distinguishing species which is a necessity for a good taxonomist. 
I, at least, have never found his systematic contributions useful 
or usable. I am not sure about the size or quality of his fern 
herbarium. Hamburg has many valuable old fern specimens, and 
many of these may have come from Prantl. 

Among the notable collections are those of Graeffe from the 
South Seas (original), Poeppig (Chile, Peru, and Cuba), Pohl 
(Brazil, probably including Presl isotypes), Philippi (Chile), Deppe 
(Mexico, isotypes of Schlechtendal), Daemel (Fiji), Sellow (Brazil, 

Ferns in European Hekbakia 61 

many Isotypes), Wawra, Zollinger (Java, Kunze isotypes), Haenke 
(Presl isotypes), Sieber (Kaulfuss isotypes), Gueinzius (South 
Africa, Kunze isotypes), and too many others to mention. One of 
the important sets is that of Karsten from Colombia and Venezuela, 
This set is so complete, the sheets so full (sometimes two and three 
sheets of the same collection), and the labels so complete ^hat I 
wonder if this is not the original set. It should be compared with 
the sets in T.eningrad and Vienna. Another notable set is from the 
botanical garden in Berlin of the early cultivated ferns described 
by link; so far as I can tell, these match closely the comparable 
specimens in Berlin, and can be considered as authentic material 

of Link's species if not actual isotypes. 

There has never been in Hamburg a pteridologist of international 
reputation except Richard Sadebeck, who was professor and cus- 
todian of the herbarium in the ISQO's and 1900's. Sadebeck had 
many interests, including ferns. Engler chose him to write the 
general account of the Pteridophyta for "Die Naturlichen Pflan- 
zenfamilien," and Sadebeck also contributed the accounts of the 
HjTnenophyllaceae, :Marsileaceae, and Salviniaceae to this work. 
Dr. C. Brick, assistant in the herbarium for many years, was also 
a fern man, and contributed the fern reports in Just's "Botanischer 
Jahresbericht" from about 1S94 to 1910. 

I visited the fern herbarium in Hamburg only briefly, for three 
days in 1954. I was cordially received by Dr. Domke, then the 
curator and was made to feel welcome. The herbarium was 
rather crowded at that time, a perennial complaint of almost all 
herbaria. I did not go to look up anything in particular but merely 
to get an idea of the scope of the fern collections. I was pleasantly 
!„„,] r .„ xx,^ v.„..u..,.;„^, ;o ;,iriocv/^ r^fli nnd should be visited 


more often by fern students. 

I am much indebted to the present curator, Dr. K 
and to the Chief Custodian, Dr. P. Wiemann, for helpful mforma- 
tion. My chief source for the historical part of this account is 
Die botanischen Institut der freien and Hansestadt Hamburg," 
by A. Voigt (1S97). 


(To he continued) 


62 American Fekn Journal 

Fertile Sporophytes of Botrychium virginianum 

Attached to Gametophytes^ 

Bruce W. INIcAlpin 

While making a fioristic survey of the pteridophytes of the 



num containing many small but fertile specimens. An apparently 
similar population has been described for the ^lountain Lake 
(Giles County) area in Virginia (Wagner, 1963, pp. 130-131.) 
Investigation of the Alacon County plants showed a considerable 
percentage of the plants attached to large gametophytes. Whereas 
juvenile plants typically are found attached to a gametophyte 
(Bierhorst, 1958), the plants found near Highlands showed a 
wide range of developmental stages, Many were fully fertile with 
small leaves 3-5 cm broad {Fig. 1), These juvenile specimens have 
had the name B. virginianum var. gracile (Pursh) Law^son applied 
to them, but are not worthy of taxonomic recognition (Weatherby, 
1935). Precociously fertile sporophytes may be due to abundant 
nutrition supplied by long-lived gametophytes. 

Gametophytes of most ferns usually die after the sporophyte 
has become self-sufficient. This is not strictly the case in the 
eusporangiate genus Botrychium where gametophytes have been 
reported still attached to sporophytes at least tw^o years old 
(Foster, 1964) and up to eight years old (Jeffrey, 1896-97, pp. 271- 
272). Campbell (1911, p. IS) mentions that fertile B, virginiamim 
sporophytes with attached gametophytes were found by Jeffrey, 
but similar plants are not mentioned by later workers. 

Bierhorst (1958) believes that a relatively small number of 
gametophytes persist after giving rise to a sporophyte. He states 
that in areas where sporophytes of all ages are present it is not the 
older sporophytes with attached gametophyte which predominate, 

* Thiti re;5earch was supported by NSF grant GB-2496 to the Highlands 
Biological Station. Voucher specimens {Wagner 69100) are on deposit in the 
herbaria of the University of Michigan and Duke University. 


but rather the very young ones. The Highlands population of B, 
virginianum that was collected in July, 1969, contained plants in 
Avhich 60% of the sterile and fertile sporophytes were attached to 
large, long-lived gametophytes. In this population gametophyte 
retention appears to be common. The attached gametophytes 
were 3-14 mm long and the largest were attached to fertile sporo- 
phytes (F?'gr, ^). 

Very large gametophj^tes could .supply abundant nutrients to 
first-year sporophytes, which then could grow larger than average 
and perhaps become fertile. The largest sporophytes had three to 
six roots and often retained the shriveled bases of previous leaves. 
Since a single root and leaf typically are produced annually by 
Botrychiiim sporophytes, this indicates that the sporophyte 
developed attached to the gametophyte for a number of years. 

Nutritional aspects of the gametophyte and its symbiotic 
fungus have not been adequately investigated. However, the pres- 
ence of stored starch in the gametophyte is evidence that the 
symbiotic process is efficient. This food reserve is presumably used 
by developing embryos and young sporelings before the sporophyte 
has become self sufficient. Conceivably the stored material and 
even the symbiotic metabohsm of the gametophyte and fungus 
may be beneficial to the sporophyte throughout its life history. 
This appears to be the case in some Botrychium plants with long- 
lived gametophytes. 


Factors Avhich cause gametophyte retention are unknown but 
appear to be frequent since most collections of Botrychium gameto- 
phytes contain individuals with attached sporophytes (Bierhorst, 
1958). In many large populations of Botrychium there is much 
variation in leaf size. A gametophyte is often attached to the 
smallest plants (leaves 1-2 cm broad). Occasionally several 
hundred sporelings with attached gametophytes can be collected 
in an area of a few square feet. Careful sifting and washing of 
the soil over a fine mesh screen may yield gametophytes in all 
stages of development. Pteridologists interested in securing 
gametophytes of Botrychium species for study or teaching material 
need only look for the smallest sporophytes in or near large popu- 

American Fern Journal 

Volume CO, Plate 7 

■_J .-L.rf/^.-_ .-/--■-■ ■ A^v^. #S- ^_- ".'_'_♦'_'---"■.■--- "-V ^' "." " 'rV ■■ "-^.-^lVx^'.V 





.'-■_*■. Pi 






Fertile BoTRvcjin'M vjrgimam m SruRupiiVTL.s with AxTACiiKD Gamk- 

TopinxEs. Fig. 1. IIauit of plant, X O.S. Figs. 2a, b. Oametophytes 


(arrow), CA. X 1. 

Peculiar Species of Gramaiitis 65 

lations. Careful search of the surrounding soil may then yield an 
entire ontogenetic series of the subterranean gametophytes. 

Literature Cited 

Campbell, D. H. 191 1. The Eusporangiatae. Carnegie last., Washington, D.C. 
BiERHORST, D. W. 1958. Observations on the gametophytes of Botrychiinn 

virginianum and Botrychium dissectum. Amer. J. Bot. 45: 1-9. 
Foster, D. B. 1964. The gametophyte of five species of Botrychium. Ph.D. 

thesis, Cornell University. 
Jeffrey, E. C. 1896-97. The gametophyte of Botrychium virginianum. 

Trans. Canad. Inst. 5: 265-294. 
Wagner, W. H., Jr. 1963. Pteridophytes of the :\Iountain Lake Area, Giles 

County, A'irginia, inchiding notes from Whitetop Mountain. Cas- 

tanea 28: 113-150. 
Weatherby, C. a. 193.). A list of varieties and fornis of the ferns of eastern 

North America. Amer. Fern J, 25 : 95-100. 



A Peculiar Species of Grammitis 


Poly podium matheivsii Kunze ex 2slett. was described from 
Peruvian specimens collected by Mathews; the locality was not 
specified but was given by Hooker (Sp. Fil. 5: 20. 1S04) as Chacha- 
poyas, and the numbers as Mathews 1811 and 3281. However, it is 
evident that :\Iathews collected this species at least three times, 
all at Chachapoyas, for the specimen in the British .Museum is 
no. 1837. The holotype in Kunze's herbarium has been destroyed. 
A specimen of one of these numbers in the Mettenius herbarium 
in Berlin should be designated as lectotype, if one exists, other- 

G6 American Feun Jouuxal 

wise one of the Kew specimens should be the lectotype. Hooker 
stated that the specimens had long been preserved in his herbarium 
as a new species. 

This peculiar and distinctive species was placed bj' Alettenius 
and by Hooker among the Goniophlebiums because of its vena- 
tion, but it is quite different in its general aspect — thick texture, 
long-decurrent segments with the lowest veinlet arising from the 
costa, and especially by the setiform hairs, which are exactly like 
those typical of many species of Grammitis. An examination of the 
spores shows that they are trilete, and that therefore this is truly 
a species of Grammitis sect. Cryptosorus. It is the only known 
species of this section with more or less completely goniophlebioid 
venation. The segments have a single, incomplete row of costal 
areoles, with one included veinlet. However, unlike the true 
Goniophlebiums, the sorus is dorsal on the included veinlet and 
not terminal. This venation pattern is an extension and compli- 
cation of the type of venation found in Grammitis kalbreyeri 
(Baker) ^Morton, comb. nov. {Polypodium kalbreyeri Baker, Trans. 
Liim. Soc. II, Bot. 2: 291. 1887) and some other large species of 
Grammitis. It is to be expected that when segments become broad 
that anastomoses can occur, a phenomenon well known within 
Polypodium sect. Polypodium (including Goniopldehium) and 
Polypodium sect. Marginaria. Thiti is not the only instance of 
anastomosing veins in Grammitis sect. Cryptosorus, either, for 
anastomoses are occasionally found in G. eminens Alorton and G. 
trifurcata, and rather uniformly in G. crispata (J. Smith) Morton. 
However, none of these species seem at all allied to Polypodium 
matheivsii, which should be known as Grammitis mathewsii 
(Kunze ex Mett.) Morton, comb. nov. {Polypodium mathewsii 
Kunze ex :\Iett. Abhandl. Senckenb. Naturf. Gesell. 2 : 74. 1856). 
The relationship of this species must remain undetermined at 
present; very hkely it is an isolated type that might constitute a 
section by itself. 

There may be some relationship to such species as Grammitis 
albidula (Baker) :\Iorton, comb. nov. {Polypodium albidulum 
Baker in Mart. Fl. Bras. 1(2): .598. 1870) and Grammitis discolor 

Peculiar Species of Grammitis C7 

(Hook.) Morton, comb. nov. {Poly podium discolor Hook. Icon. 
PL 4: t. 386, 1S41), which differ in having the blades ceraceous 
beneath, and the veins forked but not anastomosing. In (?. 
niathewsii the bhides are green beneath and not white-waxy. 

Grammitis matltcicsii is widespread; it has been known from 
Colombia to Peru, and may now be reported from Bolivia also. 
According to Hooker, Goniopfdebium villemifiianum Fee (Mem. 
Foug. 7: 63, /. 27, /. 3. 1857) is a synonym, and from Fe6's figure 
this would seem to be correct. However, Fee described and illus- 
trated some clathrate scales that are said to occur on the lower 
surface, and I have not found such scales in the material I have 
studied, nor indeed are they to be expected in this genus. F^e's 
statement may have been an error of observation. His species 
came from paramos above Ocana, Norte de Santander, Columbia^ 
at 3400-3700 m. elevation, where it had been collected by Schlim. 
I have seen a specimen from the Paramo del Hatico, Norte de 
Santander, which must not be very far away, and this agrees well 
with Fee's description and figure. 

Specimens examined: 

Colombia: Pdramo del Ilatico, Norte de Santander, alt. 290 m, Killip & 
Smith 20686. Near Facatativa, Cundinamarca, under rocks, Ariste-Josepk 
A4OS. Paramo de Guasca, above Bogota, Cundinamaraoa, Ariste-Joseph 
A344, A477. Bogotd, alt. 3,200 m, Lindig 178 (BM). Without hx^ality, Mutis 

Ecuador: Near the laguna, Paramo del Castillo, crest of the eastern Cordil- 
lera on the trail between Sevilla de Ore and ]Mendez, Azuay, epiphyte, alt. 

2,700-3,300 m, Camp E5108. 

Peru: Pendent from tree branches or rocks, frequetit, moist ravine head on 
middle eastern Calla-Calla slopes, 3-5 km. SE of km 422 on the Leimebamba 
Balsas road, Prov. Chachapoyas, Amazonas, alt. 3,000-3,200 m, Wurdack 
1765; same, near km 415^18, alt. 2,900-3,150 m, Wurdack 174S {these two 

collections from somewhere near the type locality). Alturas de Sucre, Cuzco, 

alt. 3,000 m, Bues 1538. Montana de Calea, ^'alle de Nares, Cuzco, alt. 2,250 m, 
Bues 1924. Chachapoyas, Mathews 1837 (B:\I). 

Bolivia: ^'alle de Corani, Cochabamba, alt. 800 m, epiphyte, July 6, 1968, 

Adolfo 103, 



68 A.AiERiCAX Ferx Journal 

Vascular Organization of the Rhizome of Cibotium barometz 

SuBHASH Chandra^ 

Cihotium is a genus of about a dozen species distributed in 
southeastern Asia, Hawaii, and Central America. Some of the 
species are large and rather resemble tree ferns. Cibothim barometz 
is a graceful, large, Malaysian fern with a stump-like rhizome 
5-8 cm. in diameter that appears much stouter because of its 
crowded, persistent leaf bases. Its rhizome is the ^'Vegetable Lamb" 
or "Scythian T.amb" of the Orient. The rhizome, which is densely 
clothed with large, tufted, golden bi'own hairs and hence appears 
like a wooly lamb, is semi-erect and bears a crown of large, bipin- 
nate fronds 2-3 m. long that are covered with similar hairs at 
their base. These profusely produced hairs are used for stuffing 
cushions and pillows and are reputed to staunch a bleeding wound. 

Most pteridologists believe Cibolium is cyatheoid (]\Iettenius, 
1856; Diels, 1899; Manton, 1958; Holttum and Sen, 1961; Nayar, 
1970), but others think it is related to the dennstaedtioid ferns 
(Bower, 1926; Christensen, 1938; Copeland, 1947). The anatomy 
of the Cyatheaceae, including some of the species of Cibotium^ has 
been studied recently by Holttum and Sen (1961) and by Sen 
(1964), who have pointed out the significance of rhizome anatomy 
in assessing the phylogenetic relationships of these ferns. Some 
aspects of the anatomy of C barometz were reported by Ogura 
(1926, 1930), and some details of the vascular organization were 
discussed by Hayata (1929). On the basis of the stelar structure, 
the latter separated C. barometz as a subgenus, Microcibotiiim, 

This study is based on material collected from Naphuk, Assam 



ethyl alcohol, and stained with combinations of safranin and 
either fast green, sudan IV, or iodine. 

U am indebted to Dr. B. K. Nayar, National Botanic Gardens, Lucknow, 
for his giiidance^ encouragement, and discu^itiions, and to jVh*. P. C. Roy for 
his help in making the illustrations. 

Rhizome of Cibotium Bakometz G9 

The rhizome is soft and is bounded by a thin- walled epidermis; 
20-30 layers of hypodermal cells form a sheath composed of narrow, 
slightly elongated cells having faintly thickened w alls. The ground 
tissue cells are thin-walled and parenchymatous, and contain 
dense starch deposits. Numerous cells contaniing mucilage are 
scattered in the parenchyma. The vascular cylinder {Fig. 2) is a 
large solenostele (the pith is often 4-6 cm. in diameter) pierced by 
small, subovate, spirally arranged leaf gaps {Fig. 2, g); because the 
leaves are crowded, the leaf gaps are often so closely plat'cd that 
the vascular cylinder ai)proaches the dictyostelic condition. No 
medullary vascular tissue is found. Each leaf {Fig, 2, 1) is supplied 
by a large, corrugated, gutter-shaped vascular strand given off 
from the lower end of the leaf gap. Before separation from the 
stelar cylinder, the leaf trace forms a broad, prominent, hollow, 
outward protrusion which extends some distance below the leaf 
gap. These protrusions make the cylinder irregularly corrugated. 
The leaf trace bundle, although a solitary band at its origin, 
divides into a large number of narrow strands before entering 
the leaf base. The strands are arranged in the form of an open 
gutter with incurved margins and a longitudinal, lateral invagi- 



curved margins facing the center; the other two are smaller, 
adaxial, more sharply curved arches with the concavity facing 
away from the center. The leaf trace maintains this shape as it 
passes up the stipe. The roots are mostly associated with the leaves, 
and the root traces are given off in clusters {Fig. 2, r) just before 
the leaf trace separates from the stelar cylinder. 

The xylem tissue of the vascular cylinder is com|)osed of 6-12 
layers of broad, short, scalariform tracheids interspersed with 
narrow, ribbon-Hke, interconnected bands one cell thick that are 
mostly arranged radially and composed of thin-walled xylem 
parenchyma cells. Phloem surrounds the xylem on either side as 
a thin layer composed of two or three layers (one or two only on 
the abaxial side). The phloem consists of narrow, thin-walled 
tracheids mixed Avith parenchyma cells. There is a peripheral 

American Fern Journal 

Volume 60^ Plate 8 

Anatomical Details of Cirotium barometz 


Rhtzo:\ie of Cibotiuai Barometz 71 

layer 3-5 cells broad of taiigeutially much elongated^ narrow, 
thill walled cells {Fig, 5, t) surrounding the phloem^ as is charac- 
teristic of the cyatheoid ferms (Holttum and Sen, 1901; Sen, 
1964). These cells stain deeply with fast green, are devoid of nuclei, 
and possess scattered sieve areas on all walls (Ogura, 1927; Holttum 
and Sen, 1961; Sen, 1964). This layer of tangential cells gives a 
characteristic appearance to the vascular cylinder in cross-section. 
The pericycle is two- or three-layered. The endodcrmis is poorly 
differentiated, but is demarcated from the much larger cells 
of the ground tissue. Cubical cells and sclerenchymatous bands, 
as reported in some of the Cyatheaceae (Holttum and Sen, 1961), 
are absent. The leaf trace bundle is structurally similar to the 
main vascular cylinder, but the xylem tissue is thinner and con- 
tains only 1-3 layers of tracheids; tangential cells are absent, and 
the endodermis is better differentiated. 

Vascular anatomy of the rhizome of C harometz lends support 
to the hypothesis that it is related to the cyatheoid ferns. The 
leaf trace bundle and its relationship to the main stelar cylinder 
are similar to those reported in Cyathea and related genera. The 
characteristic tangential cells surrounding the phloem tissue in C. 
harometz are paralleled in the cyatheaceous genera and constitute 
a characteristic anatomical feature of that group, but are not found 
in the dennstaedtioid ferns. Also, the vascular cylinder of C. 
harometz is radially symmetrical with spirally arranged leaf gaps, 
as in the cyatheoid ferns, in contrast to the dorsiventral stele 
of the dennstaedtioids (Kaur, 1962; Xayar and Kaur, 1963a, b), 
which have the leaf traces arranged in two dorsal rows, with leaf 
gaps lateral to them on the side facing the median plane of the 

Fig. 1. Cross-section of vascular strands at leaf base. Fig. 2. Portion 
OF stelar cylinder of the rhizome. Fig. 3. Cross-section of a portion of 

THE vascular cyllnder OF THE RHIZOME. The abbreviations are: c = cortical 



72 American Fern Journal 

Literature Cited 

Bow^R, F, O. 1926. The Fertis, vol. 2. University Press, Cambridge. 

Chrtstexsen, C. 193S. Filicinae in F. Verdoorn, Manual of Pteridologj'. "M. 

Nijhoff, The Hague. 

CoPELAND, E. B. 1947. Genera Filicum. Chronica Botanica, Waltham, Mass 

DiELS, L. 1899. Cyatheaceae in A. Engler & K. Prantl, Die Natiirlicher 

Pflanzenfamilien 1(4): 113-139. 

TTayata, B. 1929. Microoibotium, a new subgenus founded through the con- 
sideration of the stelar structure of Cibotium barometz. Bot. Mag. 
Tokyo 43: 312-317. 

IIoLTTiM, R. E. and U. Sex. 1961. ^lorphology and classification of tree ferns. 

Phytomorpholog3- 11 : 406-420. 

Kaur, S. 1962. ^lorphological and anatomical investigations on ferns: Bolbitis, 

Egenolfia and related Ferns. Ph.D. Thesis, Agra, University. 

Mantox, Irexe. 1956. Chromosomes and fern phylogeny with special refer- 
ence to *Tteridaceae." J. Linn. See. London, Bot., 56: 73-92. 

Mettexius, G. 1856. Filices Horti Botanic! Lipsiensis. L. Voss, Leipzig. 

Nayar, B. K. 1970. Scheme for a phjdogenetic classification of the homosporous 

ferns. Taxon 19 : 229-236. 

— , and S. Kaur. 1963a. Contributions to the morphology of some 

species of ^licrolepia. J. Indian Bot. Soc. 42: 225-232. 

~, and . 1963b. Ferns of India— VIII. ^Microlepia 

Presl. Bull. Nat. Bot. Gards., Lucknow 79: 1-25. 
0(;t^ra, Y. 1926. On the structure and affinity of Cibotium barometz Sm. Bot. 

:\Iag. Tokyo 40: 349-359. 

. 1927. Comparative anatomy of Japanese Cyatheaceae. J. Fac. 

Sci. Imp. Univ. Tokyo 1: 141-350. 

— . 1930. On the structure of Hawaiian Tree Fern with notes on the 

affinity of the genus Cibotium. Bot. Mag. Tokyo 44: 467-478. 
Sex, U. 1964. Importance of anatom^^ in the ph3-logen3- of tree ferns and their 

allies. Bull. Bot. Soc. Bengal 18: 26-33. 

National Botanic Gardens, Lucknow, India. 

jMorphology of Elaphoglossum 73 

A Survey of Some Morphological Features 
of the Genus Elaphoglossum in Costa Rica 

Robert M. Lloyd 


The systematics of the over 400 species of Elaphoglossum 
(Aspidiaceae) is as little known or understood as any large genus 
of tropical ferns. Although the genus as a ^vhole is pantropical, the 
majority of species are found in the Xeotropics, an area in need of 
much botanical exploration. Endemism appears to be extremely 
common. A plethora of forms occupy similar habitats in the same 
region and many occur side by side on tree trunks and branches. 
Until much additional field work is done in the Xew World tropics, 
many of the species in the genus will remain undescribed. 

Bell (1950; 19ola, b; 1955; 1956) initiated a series of mor- 
phological studies in the genus Elaphoglossum dealing with stelar 
structure, root and bud traces, and anatom^^ of the rhizome and 
frond. In these studies he made no attempt to correlate form with 
habitat. During the summer of 1967, I undertook a somewhat 
similar study in Costa Rica to add to our knowledge of morphology 
in the genus, as well as to correlate this with the basic habitat. 

Plants were collected in seven geographical locations throughout 
Costa Rica: Peninsula de Osa (0-500 m elevation), San Ramon 
(1000 m), San Vito de Java (1100-1400 m), Volcan Poas (ISOO- 
2650 m), Volcan Barba (2200 m), Cerro de la .Muerte (1200 3350 
m), and Finca La Selva, south of Puerto Viejo (100 m). The 
habitats represented by these areas vary from Pacific and Atlantic 
lowland tropical wet forest to cloud forest^ montane rain forest, 
and mossy elfin forest. Observations were made on the following 
characters: petiolar joints and swellings, pneumatophores, laminar 
scales, and sporangial features including paraphyses, number of 
indurated annular cells, and the presence or absence of chlorophyll 
in the sporangial stalk. 

The data have been grouped according to plant habitat (epi- 
phytic, epiphytic and terrestrial, terrestrial) and elevation (0- 
1000 m, lowland rain forest: 1000-2500 m, mid-elevation wet and 

74 American Fern Journal 

dry forest; and above 2500 m^ high elevation wet forest). The 
climates of these areas are discussed by Scott (1966). 

The data presented here represent only a summary of that 
collected. Mimeographed sheets with full results will be provided 
by the author on request. Voucher specimens have been deposited 
in the herbarium of the New York Botanical Garden. About 80 
species were investigated, 70 in detail. Some of these are un- 
described and the relationships of many others are in doubt. 

Table I. Habitat of ELAPiiO(iLossi:M by Elevation* 

Numher of species and percent 
epi~ terr. <& ter res- 

Elevation phytic % epi. % frjal % 

0-1000 m 13 93 1 7 

1000-2.500 m 29 76 9 24 

above 2500 m 14 50 4 14 10 36 

0-2o00m 42 81 10 19 

ToUl 56 70 4 5 20 25 


Habitat. ~~0i the 80 species of Elaphoglossum studied, 56 (70%) 
were epiphytic. There is a direct correlation between increasing 
elevation and frequency of terrestrial habit. At elevations below 
1000 m, over 90% of the species were epiphytic, but above 2500 m 
only 50% occupied this habitat (Table I). In addition, those 
occupying both terrestrial and epiphytic habitats were found only 
at higher elevations. 

Petiole Characters. — There is di^tini^t variation in the petioles 
in Elaphoejhsswn. At the base of the petiole in the region where 
abscission occurs there may occur a joint or swelling. The joint 
refers to a point where there is a distinct color change where the 
petiole and phyllopodium join. This may or may not be accom- 

^loiiPHOLOGY OF Elaphoglossum 75 

pamcd by a slight swelling of the region (see Bell, 1955, fiy. 4.). Of 
the 70 species investigated, 59 exhibited a distinct color change 
from petiole (usually green) to phyllopodium (usually dark brown 
or black). In 54 of these the joint was accompanied by swelling. 
Only five species possessed joints without swelling: E. cxiinium 
(Mett.) Christ, E. aff. lindenii (Bory) Moore, E. muscosum 
(Swartz) Moore, Lhijd 4£19, and Lloyd 4220. The eleven species 
which were unjointed did not possess swellings. These results are 
similar to those of Bell (1955), who found a dose association 
between the two characters with but only one exception. 

Terrestrial species exhibited a greater frequency of joints and 
swellings than epiphytic {Table II), and those from middle eleva- 
tions appeared to have a greater frequency with both structures 
than those from either lowland or highland areas (Table IV). 

Pneu7natophores. — Aerenchymatous flanges (or piieumatophores) 
are composed of loose, parenchymatous cells with large air spaces; 
they usually extend as a flange through the sclerified outer cortex 
of the phyllopodium. Bell (1955, fuj. 4) recognized several types. 
They vary from narrow, lenticel-like lines extending from the base 
of the phyllopodium to the petiole to elaborate wing-like or phal- 
loid structures of various sizes. They have been found in 92% of 
the species in Costa Rica, with the most frequent types being 
•either a wavy, linear and somewhat fleshy tissue about 1-2 nmi 
broad extending the length of the phyllopodium (Type C), found 

hhtiim (Swartz) C. Chr., E. hyalinum Christ, E, aff 




long and 1-2 mm wide, arising from and restricted to the region 
near the base of the phyllopodium (Type G). This type of pneu- 
matophore is always associated with densely seal}' rhizomes and 
IS found HI 1S% of the species, including E. conspersurn Christ, E. 
hermimeri (Bory & Fee) :\Ioore, E. lonyifolium (Jacq.) J. Smith, 
E, revolvens (Kunze) C. Chr., and E. squamipes (Hook.) Moore. 
The most common type of pneumatophore (C) occurs hi 46% of 
the epiphytic species and onlj^ 15 %. of the terrestrial. Type G is 

76 American Fern Jourxal 

Table II. Percent of Species with Characteristics in 

Relation to Epiphytic, Terrestrial and Epiphytic, 

AND Terrestrial Habitat 

Chamder epiphnlic ten. i^ cyi. krrcslrial Mean 

Swelling 7S 50 85 71 

Jointed 80 75 95 SO 


Typo A 4 20 9 

B 12 5 S 

C 46 75 15 30 

1) 4 15 7 

E 6 10 7 

F 13 10 11 

G 15 25 25 IS 

Sr-ales: adaxial 

Glabrous 50 25 00 52 

Type A {fig. 8-1 y> 2 1 

B (Jg. 7) 14 25 5 12 

C {fig. 8-7) 14 25 30 li) 

D {fig. 8 10) 7 4 

E{fg.8-ll) 13 25 5 12 

Coverage : 

Under 10% 80 25 80 77 

10-40% 15 75 15 19 

Over 40% 5 5 4 

Scales: abaxial 

Glabrous 14 20 15 

Tyjw A 2 1 

B 9 10 9 

C 52 25 50 50 

D 7 50 5 9 

E 16 25 15 10 


Under 10% 80 50 85 79 

1040% 2 2 

Over 40% IS 50 15 19 



present 6S 43 56 

absent 32 57 44 


found in 15% of epiphytic forms and 25% of terrestrial. There is 
a trend to lack of pneumatophore in terrestrial species, with 20% 
lacking this structure. In the epiphytes only 4% were lacking it. 

Mature pneumatophores were frequently observed on the 
rhizome in areas of leaf primordia. In many instances as the 
petiole developed, the tissues of the pneumatophore appeared to 
fuse with those of the petiole. The function of the structure is 
unknown, although Troll (1933) believed that it was for aeration 
in areas of high metabolic rate, especially where aeration was 
restricted, such as croziers with heavy mucilage. This substance 
has not been found in Elaphoglosswn, however. 

Although Type C is the most common type of pneumatophore, 
at elevations above 2500 m there is a sharp increase in number of 
species without pneumotophores and with Type G {Table IV). 

Laminar Scales, — The structure of scales and hairs on the frond 
varies from simple or capitate glandular hairs to peltate scales. 
The variation in structure found in plants from Costa Rica is 
similar to that found by Bell (1955, fig. 8) for plants from other 
areas of the world. The most common scale type was laminate, 
attached at the base with cordate or lobed basal wings (Belly fig. 8- 
7), In only six species did scale types differ on the adaxial and 
abaxial surfaces: E. auripilum Christ, E. aff, hirtum (Swartz) C. 
Chr., E. muscosum (Swartz) Moore, E. revolvens (Kunze) C, Chr., 
E. aff. tectum (Humb. & Bonpl.) Aloore, and Lloyd 4220. In E, 
auripilum and E. aff. tectum the more complex scale type was 
found on the adaxial surfaces. 

Over half of the 70 species examined in detail possessed no 
scales at all on the adaxial surface. Of those with scales, 77 %» had 
a coverage of less than 10 % and only 4 % had a coverage of greater 
than 40%) {Table II). Sixty percent of the terrestrial species and 
50%) of the epiphytic were glabrous. In most instances the trends 

Pneumatophore typeri: A. None. B. Lenticel line extending from the base 
or the phyllopodium into the petiole; no flange. C. Lenticel line present with 
slight flange near the base. D. No lenticel line; slight flange present at ba^e of 
phylU)pudium. E. No lenticel line; definite flange present. F. No lenticel hue; 
enlarged flange, G. No lenticel line; club-shaped flange, 
^ Figures of the various t3-pes are from Bel! (1955). 


American Fern Journal 

present on the adaxial surface were similar to those found on the 
abaxial surface. On the abaxial surface, however, scale density 
was greater, with about 18% of the species having over 40% 

Elevation apparently has no profound effect on either scale 
type or density. From the data given in Table III, there appears 
to be only a slight trend toward more scaliness on the abaxial 


Table III. Number of Species of Each Scale Type 

IN Relation to Elevation 

Scale type'' 












Adaxial Surface 


0-1 000 m 




1000-2.-) 00 m 








Above 2.300 m 







Abaxial Surface 

0-1000 m 




1 000-2500 m 








Above 2.-)00 m 







* See Table II for scale types and references to illustrations in Bell, laV). 
^ Based on 13 species, 4 of which were analyzed as to type. 

surfaces at higher elevations. However, this trend appears to be 
reversed on the adaxial surface. 

Para/7^?/se.s.— Copeland (1947, p. 119) stated that paraphyses 
are wanting in Elaphoglossum. However, recent studies by Ander- 
son and Crosby (1966) indicate the presence of paraphyses in 
some Hawaiian species. A survey of fertile fronds in Costa Ilican 
material indicates that 56 % have paraphyses. In most cases the 
paraphyses are short, uniscriate, multicellular branches arising 
from the upper portion of the sporangial stalk near the base of 
the sporangium. In a few cases they arise from the receptacle 

^NFoKPHOLOGY OF Elaphoglossum . 79 

{Lloyd 4268) or from both the receptacle and the sporangial stalk 
(Lloyd 4281,). They are usually glandular at maturity. Paraphj-scs 
occur in CS7o of epiphytic species and only 43% of terrestrial 

{Table II). 

Table IV. Percent of Species with Characteristics 

I.N Relation to Elevation 

Character 0-1000 m 10OO-2."j00 »i 2.")00-a6ot;e 

Swelling Present 75 84 6S 

Jointed 75 90 79 

Type A" 25 5 H 

B 13 4 

C 45 36 


8 7 

25 5 7 

25 13 7 


G 25 11 28 

Paraphyses Presentt- 100 54 61 

No. species investigated 4 38 ^o 

"See Table II for description of types. 

*> Based on percent of taxa with fertile fronds. 

Asexual Reproduction.— YegeiaWve reproduction is infrequent 


material it was found in only one species, E. undulatum (Willd.) 
Moore {Lloyd 4177). Plants of E. undulatum occur in very wet 
secondary forest near San Ramon and regularly produce asexual 
buds from the terminal portion of the midvein of the frond. This 
species was usually epiphytic, normally between 0.5 and 1 meter 
above ground level, with pendent fronds which frequently came 
in contact with the soil. Buds and new plantlets were observed 
on mature fronds, increasing in size as the parent fronds aged. 
As the parent fronds degenerate, the new plantlets formed ter- 
restrial colonies at the base of the tree. 

so American Fern Journal 

Other Sporangial Features. — Sporangia tend to arise in numerous, 
circular sori below the veins. As the fertile fronds mature, these 
sori merge together and become acrostichoid. The number of 
indurated annular cells of the sporangia vary from 10 to 15, with 
12 being the most frequent. Chlorophyll is present in the sporangial 
stalk in about one-third of the species. 


The most obvious trend found in this study in Elaphoglossum 
is toward the terrestrial habitat with increasing elevation. In 
lowland rain forest, 93 % of the species were epiphytic. In highland 
wet forest (above 2500 m) only 50% of the species were ejMphytic. 
This trend is frequent in other groups of plants, including Bryo- 
phyta, Bromeliaceae, and Orchidaceae. At lower elevations species 
are either terrestrial or epiphytic, but not both. At upper eleva- 
tions, particularly in exposed habitats, this distinction breaks 
down. The cause of this phenomenon is only vaguely known. Wet 
cloud forest, with its somewhat stunted trees, has altered the 
terrestrial habitat and permits normally epiphytic species to 
grow. This may be due to increased humus, mosses, water and 
light. At lower elevations the available moisture in the soil is 
greater than that on the trees, but here other factors, such as lack 
of light or available substrate, must control plant distribution. 
There is little evidence to indicate whether the terrestrial habitat 
is less extreme than the epiphytic, although it is apparent that the 
former may provide the more suitable habitat during dry periods 
in those areas with a marked wet-dry climate. Terrestrial habitats 
also provide greater protection from drying winds. Some terrestrial 
habitats, especially those of disturbed areas along roadsides where 
there is no protective cover, appear to have more extreme condi- 
tions. Species of Elaphoyhssum that occur in these habitats have 
coriaceous fronds, a feature that Lloyd (1965) found to be as- 
sociated with xeric environments in Pohjpodiwn. 


Morphology of Elaphoglossum 81 

There are certain characters correlated with either epiphytic or 
terrestrial habitat. These are: 

Epiphytic TerreMrial 

Non-jointed Jointed 

Non-swelling Swelling 

Type C pneumatophore Type G pneumatophore 

Pneumatophores present Pneumatophores absent 

Scaly Glabrous 

Type B, C, & E scales Type C scales 

(adaxial surface) (ad axial surface) 

Paiaphysate Non-paraphysate 

The strongest of these trends is toward the absence of para- 
physes in terrestrial plants. In addition, there is a shght increase 
in the frequency of paraphyses with higher elevations, although 
the number of terrestrial species also increases. If paraphyses 
evolved as protective structures against desiccation of young 
sporangia, epiphytic plants more exposed to changing conditions 
would be expected to show a higher frequency of presence. This 
might also be true of plants in more extreme situations at higher 

Bell (1955) found a close association between the joint and the 
presence of pneumatophores, although he beheved that they 
were probably independently controlled genetically. The function 
of the pneumatophore is unknown. The function of the joint may 
be as a mechanism for shedding leaves, as in Oleandra. Holttum 
(1966) believed that this was an adaptation to the epiphytic 
habit. However, in Elaphoghssum , terrestrial plants show a higher 
frequency with joints than do epiphytes. Species at higher eleva- 
tions also possess this structure less frequently. 

The trends in scaliness are of uncertain significance. Plants 
"v^^th both glabrous and densely scaly fronds occur side by side in 
many different habitats. Christ (1899) believed that species at 
higher elevations tended to have densely scaly fronds and those 
at lower elevations glabrous fronds. JNIy results indicate that the 
adaxial surface of the frond is less densely scaly at higher eleva- 
tions than at mid-elevations. However, on the abaxial surface 

82 American Fehn Journal 

scale density increases with elevation. Over 50% of the species 
from elevations above 2500 m had glabrous fronds or fronds with 
less than ten percent of the surface covered by scales. 

It is obvious from this study that the relationship between the 
characters investigated and the habitat is complex. The wide 
diversity hi structure may be due to maintenance of much of the 
diversification through time since the origin of the genus in a 
stable, continuously tropical habitat (Bell, 1956), and in fact, 
there may be no correlation with habitat for many of the features 
I studied. Further studies are needed, however, before definite 
conclusions can be reached as to the evolution of the various 
features and their relationship to the environment. 

Use of Characters for Taxonomic Purposes. — Many of the fea- 
tures elaborated in this study appear to be useful for taxonomic 
delineation of species. Of particular importance are scale type, 
pneumatophores, presence or absence of a joint and phyllopodium, 
and presence and type of paraphyses. Bell (1955) found that the 
scale types are constant in any one species. This may be true of the 
other features, although large numbers of mdividuals of each 
species have not been investigated. Nearly all 70 species can be 
distinguished using a combination of these characters. 


Much of this work was done while I was involved in a course on 
the biology of tropical pteridophytes, sponsored by the Organiza- 
tion for Tropical Studies. I am indebted to that organization and 
to the University of Costa Rica for providing funds and facilities 
while in Costa Rica. I am also indebted to Drs. W. H. Wagner, 
Jr., J. T. Mickel, D. B. Lellinger, A. M. Evans, and C. V. Morton 
for their helpful criticisms and comments during the study. Speci- 
men identification was done by Drs. Mickel and Lellinger. 

Literature Cited 

Anderson, W. R. and M. R. Crosby. 1966. A revision of the Hawaiian speriew 

of Elaphoglospum. Brittonia 18: 380-397. 
Bell, P. R. 19r)0. Studies in the genus Elaphoglossum Schott. I. Stelar strvu- 

ture in relation to habit. Ann. Bot,, Lond. n.s. 14: r>4.')-5r)o. 

Notes and News 83 

— . 1951a. Stadias in the genius Elaphoglossnm. II. The root and 
bud traces. Ann. Bot., Lond. n.s. 15: 333-346. 

19r)lb. Studies in the genus Elaphoglos^^um. IIL Anatomy of 

the rhizome and frond. Ann. Hot., Lond. n.s. 15: 347-357. 
— .1955. Stiidies in the genus Elaphoglopsum. IV. The morphological 
series in the genus and their phylogenetic interpretation. Ann. Bot., 
Lond. n.s. 19: 173-199, 

— — . 1956. Studies in the genus Elaphoglossum. I\'. Part Two. Ann. 

But., Loud. n.s. 20: G9-88. 
Christ, II. 1899. JVIonographie des genu.^ Elaphoglossum. Denkschr. Schweiz. 

Naturforsch. Ges. 36(1) : 1-159. 
CoPELAND, E. B. 1947. Genera Filicum. Waltham, ^ilass. 
IIOLTTL'M, R. E. 1966. Florae ^[alesianae Precursores XLI\'. The fern gemis 

Elaphoglossum iji Malaysia, with descriptions of new species. Blumea 


Lloyd, R. M. 1965. Clinal patterns in frond anatomy of Polvpodium. ]\Iadn»no 

18 : 65-74. 

Scott, N, J. 1966. Ecologically impoi'tant aspects of the climates of Costa 

Rica. Organization for Tropical Studies, Mimeo. 

Troll, W. 1933. Botanische Mitteilungen aus den Tropen. IX. Dryopteris 

sumatrana v.A.v.R., ein neuer Schleimfarn aus dem astindischen 
Archipel, Flora (Jena) n.f. 28: 329-337. 

Departmext of Botany, Univeksity of ILiwaii, Honolulu, 

News and Notes 

Fern Foray Participants should ^vrite for details to Miss Lois 
Carlson, Alatthaei Botanical Garden, University of Michigan, Ann 
Arbor, :\Iich. 48105 if they have not already done so. 

Mrs. Charles Bittinger, who was instrumental in developing 
the Fern Valley at the National Arboretum in Washington, D. C. 
during the I950's, died in Washington on April 24 at the age of 95. 
She had been active in garden club activities in the Washington 
area for many years, and had received honors from the Garden 
Club of America, She was also interested in music and the other 
^rts, and was a member of the Women Geographers, an arm of the 
Explorer's Club. She described the establishment and growth of 
the Fern Valley in a well illustrated article several years ago 
(Amer. Fern J. 51: 161-178. 19G1). 

84 American Fern Journal 

Recent Fern Literature 

The Victorian Fern Craze, by D. E. Allen. Hutchinson and 
Co., Ltd. 178-202 Great Portland Street, London W. 1, England. 
$3.00 postpaid. 83 pp. illus. 1969.— Today's growers of ferns are, 
by and large, a sensible lot, who grow their ferns in plastic pots 
on window sills and who even may have some scientific interests 
that run to the detection of hybrid ferns or the discovery of the 
life cycle of some particularly interesting species. A century ago. 
however, such was not the case. Gardeners, particularly in Great 
Britain, were seized with a pteridomania that rivaled the tulip 
mania which had seized Holland some time before. Hybridization 
of ferns being difficult and full of chance, the fern fanciers of 
Victorian England combed the wilds of their island in search of 
sports and fancy varieties of their native ferns. In many cases, 
they decimated natural populations. They searched their gardens 
and greenhouses for specimens varying in texture, dissection, 
and habit, all with an eye to presenting a striking novelty at the 
next garden show. The principal stimulus to this frantic activity 
was the popularization of Wardian cases which, being closely 
glazed, developed a high humidity suitable to the growth of ferns. 
Even filmy ferns could be maintained in an average home in a 
Wardian case. Unlike the simple terraria or bell- jars of today, the 
closely glazed cases of a century ago were ornamented with con- 
siderable quantities of intricately carved woodwork. They formed 
a large and very fashionable part of the furnishings in the best 
homes of the time. The discoveries of Dr. Ward and others, the 
production of books on ferns by both amateur and professional 
botanists, the pteridomania itself, and the eventual slackening of 
interest in fancy ferns are all admirably told by Mr. Allen in this 

delightful book.— D. B. L. 

Welsh Ferns, Clubmosses, Quillworts, and Horsetails, 
A Descriptive Handbook. By H. A. Hyde and A. E. Wade, 
ed. 5, by S. G. Harrison, xii + 178 pp. illus. 1969.— Despite the 
title of "Welsh Ferns" this useful handbook treats all the native 

Recent Fehx Literature 85 

ferns of the British lales. This new edition is thus the most recent 
treatment of the ferns of this area. Mr. Harrison has taken ac- 
count of current work on hybrids, chromosome numbers, and 
nomenclature, and has added a completely new section on the 
fern alheS; which were not discussed in the previous editions. This 
new edition is thus useful even for those having an earlier edition. 
A review of this new edition was written by C. Jermy (British 
Fern Gazette 10: 105, 106. 1969), in which a few nomenclatural 
changes were suggested. This new edition not only divides the 
Polypodiaceae into numerous smaller families but even into several 
orders. It may be mentioned that two of these family names are 
incorrect, Gymnogrammaceae and Aspidiacoae, since they are 
based on illegitimate generic names. I want to mention only one 
nomenclatural point, a rather troublesome one. So long as Poly- 
podium camhricum Ij. is considered as a variety of P. vulgare L. 
there is no problem. However, when P. vulgare is split into three 
species, as here, with different chromosome numbers, the matter of 
priority enters. True P. vulgare is a tetraploid, and the diploid is 
now called P. australe F^e (1852). Miss Shivas has found that P. 
camhricum is also diploid, and so in this work it is placed as Poly- 
podium australe 'Cambricum'. This form would seem to indicate 
that it is a cultivar, but it is not, for it was described from wikl 
material and is still found in the wild. The nomenclatural point is 
that P. camhricum L., dating from 1753, antedates P. australe Fee 
by a hundred years and is thus the correct name for the species. 
The reason that it has not been adopted is probably because it 
represents an abnormally cut form. One might argue that Art. 7i 
of the Code could be applied and the epithet camhricum rejected as 
being based on a monstrosity, which means that it can not be used 
at all, certainly not as a variety of P. australe. However, m my 
opinion, Art. 71 was not intended to make names of this sort 
illegitimate; that would mean that no abnormal forms could ever 
be named, for these names would automatically be illegitimate 
and rejected. A solution to this dilemma in this particular case is 
to regard P. camhricum not as a variety of anything but as a 
possible hybrid P, X camhricum. I do not know if this has ever 

SG American Fekn Journal 

been suggested, but it is completely reasonable and probably 
can not be disproved. The plants do have ''hybrid irregularity" and 
are completely sterile. Since it is a diploid, one of the parents would 
be P. australe; it is impossible to say what the other parent might 
be, since there is no other diploid Poly podium in the area or indeed 
in all of Europe. Still, this assumption would be sufficient to 
maintain the name and P, australe also and therefore it should 


probably be made in order to avoid a confusing and somewhat 
misleading change of name for a common species. — C.V.^I. 

Fekn Hybridizing at tfie University of Leeds, by John D. 
I.ovis. Reprinted from the British I^eru Gazette, British Museum 
(Natural History), I^ondon S. W. 7, England. $0.50 postpaid. 
8 pp. 196S. — Dr. Lovis has written an interesting and detailed 
review of the hybridization techniques that have been used so 
successfully by pteridologists trained at the University of Leeds. 
He gives some useful methods for gathering and sowing spores, 
preventing contamination of cultures by unwanted spores and 
various extraneous organisms, for transplanting minute prothalha, 
and for producing cultures ready for hybridization at the proper 
time. An exacting but rather foolproof method of hybridizing is 
also presented.— D. B. L. 


Three Species in Aspleniu.u, by S. Tatuno and S. Kawakami. 
Bot. Mag. Tokyo 82: 436-444. 1969.— The three species studied, 
Aspleniu7n incisum Thunb., A. oligopJilebium Baker, and A. 
tripteropm Xakai, are all normal diploids, with 2n = 72. The 
authors have studied the chromosome morphology and can 
divide the chromosomes into six types, each with two subtypes. 
From these observations they conclude that the true base number 
for Asplenium is 12, and that these "diploid" species are really 
hexaploids. I beUeve that this is the first demonstration in support 
of the belief that modern ferns with high chromosome numbers 
are really polyploids of ancient ancestors with low numbers. 

Recent Fern Literature S7 

The Pteridophvta of Tai\van-5. Hvmenophvllaceae: Tri- 
chomanes, by Charles E. DeVol. Taiwania 14: 89-132., t. I-I4. 
1968.— DeVol has continued his vahiable series of papers on the 
ferns of Tai\A-an with an extensive study of Trichomanes, in which 
a special emphasis is given in the descriptions and illustrations to 
the cellular structure of the blades, very likely an important 
character in this genus and one that is msufficiently known. Van 
den Bosch, the monographer of the genus, paid attention to this 
character, sometimes to the relative neglect of other possibly more 
usable ones, but he has been little followed. Although DeVol uses 
the name I'richomanes in his title and in the title to the key to the 
species, he actually does not recognize the genus in the inclusive 
sense but instead recognizes all of Copeland's small segregates, 
indicating that they are 'Svell known and easily recognized 
groups," a. conclusion that I am unable to agree with. He also 
does not accept my typification of Trichomanes on T. scandeus L., 
without disputing the argument but for the reason that this species 
belongs to a small section that has fronds with ciliate margins and 
comes from the West Indies, but the size of a group that the type 
species belongs to is strictly irrelevant to its typification, as are 
also its geographic provenience and characters. Tn my discussion 
of the typification of Trichomanes (Contr. U. S, Xat. Herb. 38; 17 
1968) I mentioned that Dr. Holttum had questioned Copeland's 
typification without making any change himself, but I overlooked 
a previous paper by Holttum (Gard. Bull. Str. Settl. 12: 304. 
1940) in which Holttum did go into the matter and came up with 
the same conclusion as I, namely that John Smith's selection of 
T. scandens as lectotype in 1875 can not be altered. That really 
should not cause any inconvenience, because this was the usual 
view up until the time of Copeland's 1938 paper.— C.V.M. 


SS American Fekn Journal 

American Fern Society 

New Members 

Mr. Abdul Samat Din Abdullah, School of Biul. Sciences, Univ. of Malaya, 

Kuala Lumpur, Malaysia 
Mr. Ivan A. Butcher, Dept. of Biol. Sciences, Portsmouth Polytechnic, Hay 

Street, Portsmouth, Hants., England 
Miss Wilma A. Converse, 4803 46th Ave. S.W., Seattle, WA 98116 
Mr. M. D. Dimitrievich, 549 Rivard Blvd., Crosse Pointe, INII 4S2;]0 
Mr. Lionel N. Eleuterius, Gulf Coast Res. Lab., P. O. Box AC, Ocean Springs, 

i\II 39564 

Mr. Glenn H. Haiti, Los Angeles State k County Arboretmn, Box GSS, 

Arcadia, CA 910U6 
Mr. Johs Ilovland, 5610 Oystese, Hardanger, Norway 
Mr. Leo D. Leichter, R. D. 1, Wind Ridge, PA 15380 
Airs. Clare Booth Luce, 4559 Kahala Ave., Honolulu, HI 9G.S1G 
Mrs. R. Alan McMillan, 203 Rankin Dr., Creeneville, TN 37713 
Mr. Lawrence Plot nick, 249-05 61st Ave., Little Neck, NY 11362 
Dr. A. Rashid, Dept. of Botany, Univ. of Delhi, Delhi 7, India 
Mr. Alfred J. Roth, R. R. 1, Box 246A, Sebastian, FL 32958 
Miss :Mary J. Swanson, 314 McCalmont Hall, Menomonie, WI 54751 
Mr. William B. Weglicki, 126 Sewall Ave., Brookline, MA 02146 
:\rr. Thomas Zanoni, R.P.O. 6006, Rtttgers— The State University, New 

BrunswJck, NJ 08903 

Changes of Address 

Mr. Joel 11. Beasley, 2244 A'irginia Place N.E., Atlanta, GA 30305 
Mr. Eugene A. Bond, West Road, Dorset, VT 05251 
Mrs. Stewart P. Coleman, 6 Shadowlawn St., Houston, TX 77005 
Mr. G. D. de Joncheere, M. Y. Olie-Scheepvaart, P.O. Box 846, Rotterdam, 


Mrs. Katheryn Eastman, Kader Sokata 8/3, (;a7i Osman Pasa, Ankara, 


Mr. Samuel C. Johnson, c/o A. A. A., Box 2324, Houston, TX 77001 

Dr. C. Don :MacNem, Div. of Natural Science, Oakland Museum, lOtli & 

Fallon Sts., Oakland, CA 94612 
Dr. James P. Poole, 10 E. Wheelock St., Hanover, Nil 0;j75o 
Mr. William F. Reeve, Ontario Dept. of Mines, Kenora, Ontario, Canada 
Mrs. Marion B. Rhodes, Rua.sen Road, Sunderland, MA 0137.', 
Mrs. Laurence E. Skog, Bailey Hortoriuni, 467 Mann Library Cornell Univ., 

Ithaca, NY, 14850 
Mr. John Whitehead, :i37r, Ilualapai Road, Kingman, AZ S()401 

bntk m4 Rarif Ftrii 



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St. Helena, Califomia 94574 

Saturday* and Sundays from 10 A.M. to 4 P.M. and by appointment 

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Mall arders acccptt d 


Vol. 60 July-September, 1970 No. 3 

American jFern journal 


Published by tbe 









Stelar Anatomy of Six Species of Lycopodium. .Frances M. Cardillo 89 

A Note on the Gametophytes of Cystopteris tentiisecta S. S. Bir 98 

The Lectotype of Polypodium leptophyllum L C. V. Morton 101 ' 

Taxonomic Notes on Ferns, IV C. V. IMorton 103 

A Reinvestigation of the Morphology of Hypodematium crenatixm 

B. K, Nayar and Nisha Bajpai 107 

Shorter Note : Pyrrosia princeps, a Fern New to Cultivation. 118 

Recent Fern Literature 


American Fern Society M"?.^.??!'.^ . ^?Tf .'I'f^.'!^ ... 128 

OCT 1 2 1970 

2C()e glmerican Jfern g»acietp 

Council for 1970 

Warren H. Wagner, Jr., Department of Botany, University of Michigan, 
Ann Arbor, Michigan 48104. President 

John T. Mickel, New York Botanical Garden, Bronx Park, Bronx, New 
York. 10458. Vice-President 

A, Murray Evans, Department of Botany, University of Tennessee, Knox- 
ville, Tenn. 37916. Secretary 

LeRoy K. Henry, Division of Plants, Carnegie Museum, Pittsburgh, Penn- 
sylvania 15213. ^ ^ ^ Treasurer 

David B. Lellinger, Smithsonian Institution, Washington, D. C. 20560. 


Warren H, Wagner, Jr., University of Michigan A.AA.S. Representative 
RoLLA M, Tryon, Jr., Harvard University AJ.B.S. Representative 

glmcrican Jfern 3Fournal 


David B. Lellinger Smithsonian Institution, Washington, D. C. 20560. 

C. V. Morton Smithsonian Institution, Washington, D. C. 20560. 

RoLLA M. Tryon, Jr. 

Gray Herbarium, Harvard University, Cambridge, Ma^. 0213S 

Ira L, Wiggins Dudley Herbarium, Stanford University, Stanford, Calif. 


An illustrated quarterly devoted to the general study of ferns, owned by 
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Matter for publication should be addressed to the Editor-in-Chief. 

Changes of address, applications for membership, subscriptions, orders 
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American Jfern Journal 

_ im M_i^^ I M ii^^iiii "■ ^ 

Vol. 60 July-September, 1970 No. 3 

Stelar Anatomy of Six Species of Ly copo dium^ 

Sr. Frances ]\I. Cardillo 

A review of the literatui-e indicates that only limited studies 
have been made on the stelar patterns of the club mosses and that 
little has been attempted in comparative investigations. HoUoway 
(1916) found that the vascular cylinders of New Zealand lycopods 
exhibited specific stelar patterns. He thought that the cortical 
sclerenchyma atTected stelar arrangement and that the presence 
of this tissue could be responsible for imparting rigidity and 
lateral pressures to the stele. This rigidity and lateral pressure 
support the stele in such a way that the xylem elements become 
more definitely arranged in groups or bands. 

Later, HoUoway (1919) concluded that there were two basic 
patterns in which the xylem elements were arranged; these pat- 




in which the vascular cylinder becomes strongly developed and 
the leaf traces afiPx themselves to it. He interpreted the banded 
type as a specialization of a radial pattern that was caused by a 
restriction on the stem ui monopodial branching. Holloway 


prothalli were characteristic of the mixed type of vascular cylinder 
and long-lived prothalli of the banded pattern. He noted that 
these differences were exhibited in all the species of Lycopodium 

• This study represents a portiun of a doctoral dissertation subniittwl to 
the Graduate" School of St. Bonaventure University;, St. Bonaventure W. Y. 
Appreciation is extended to Dr. Alfred F. Finocchio for advice during the 

course of the research. 


American Fern Journal 

Volume 60, Plate 9 



P. PAR: 


Tracings of Composite Photographs of the Vascular Cylinders of 
Lycopodium lucidulum and L. 

metaphloem, m.x. = 
PROTOPHLOEM, and p.x. 

ANNOTiNUM. The abbreviations are: m.p 
metaxylem, p. par. = phloem parenchyma, p.p 


AnatojMY of Lycopodium 91 

which he studied, regardless of whether their growth habit was 
vertical or horizontal. 

This study was undertaken to examine additional species of 
Lycopbdium, A comparative investigation of the transverse 
sections of stems of six species found in the northeastern United 
States revealed similar stelar patterns in addition to several other 
observations. The species collected in southern Cattaraugus 
County, New York (Cardillo, 1967) were: Lycopodium lucidulum 
Michx., L. flabelliforme (Fernald) Blan chard, L. tristachyuni Pursh, 
L. davatum L., L. ayviotinum L. var. afmotimim, and L. obscurum L. 

First, stem sections of the same species taken at various distances 
from the apex were compared. These showed slight differences 
which could be attributed to age (Bower, 1935). It was evident, 
however, that such differences did not alter the basic stelar pattern. 
For comparative study transverse sections were taken from sub- 
apical regions of the main stems where maturation was kno^ii to 
have occurred. This would rule out the minimal differences in 
stelar patterns attributed to age. 

Tissues were killed and fixed by freeze substitution and de- 
hydrated in cold absolute ethyl alcohol (Jensen, 1962). Iodine- 
potassium iodide served to detect phloem and phloem parenchyma; 
phloroglucinol-HCl was used to detect xylem (Roberts & Herty, 
1934). Since this investigation dealt with a comparative study of 
mature stelar patterns and the cortical sclerenchyma was not 
under study except for its effect on supporthig vascular tissue in 
the living plant, the cortical material was removed just before 
killing and fixing. Thus, steles were processed instead of entire 
stem sections. This facihtated histological procedures and gave 
more accurate results. Fisher Tissuemat (melting pt. 56.5'' C) 
was used for embedding. Sections were stained with basic fuchsin 
and counterstained w^ith fast green so that a cellular inclusion 
;ivnig a positive Feulgen reaction could then be identified as a 

nucleus (Jensen, 1962). 

Since cell diameters ranged from 3 m to 50 /x, it was necessary to 
examine each transverse section at a magnification of 430 X or 
higher. In order to make a comparative study of cellular size and 

American Fern Journal 

Volume 60, Plate 10 


100 M 

Tracing of a Composite Photograph of the Vascular Cylinder of 
Lycopodium obscurum. For abbreviatioas see Plate 9. 

Anatomy of Lycopodiu.m 93 

shape, illustrations of these sections were made. Figs. 1-6 are 
accurate drawings that were traced from composite photographs 
of each stele. Such composite photographs were compiled so that 
the details of very small cells would not be lost. 

Differences not previously reported were revealed in this study. 
The results agree with Wardlaw's (1965) statement that dif- 
ferences of detail are only exposed in a comparative study of species 
in the same genus. Table I gives a comparative description of 
these details. These results also substantiate Bower's (1935) 
description of the stele of Lycopodium in that the xylem tracts 
are not distinct from one another for any great distance along the 
stem. The xylem structure anastomoses and there is a continuous 
invagination of the phloem and conjunctive parenchyma. In some 
areas of the plant, the stele with respect to stem diameter enlarges, 
but a relatively constant surface proportion of living tissue to 
adjoining dead tracheids is always maintained. 

"Very few nuclei were detected in any one section. ^^lacerated 
preparations and longitudinal sections of the stele showed sieve 
cells and phloem parenchyma cells of uimsual length. These cells 
also exhibit long and considerably vacuolated nuclei. Whether or 
not cells containing these nuclei were protophloem, phloem 
parenchyma, or pericycle could not be determined. No attempt was 
made to identify these cells in macerated preparations; in longi- 
tudinal sections the only way to identify the cell type is by position 
in respect to the xylem and the sieve areas located on the walls of 
the phloem and protophloem. Cells of the central portion of each 
phloem band were enucleate, but nuclei were detected in cells on 
the periphery of these bands. In longitudinal section, these cells 
of the central portion appear thin walled and are marked with 
sieve areas. Cells of the protophloem located at the periphery of 
the stele and intermitent vnih protoxylem bear similar markmgs. 
These cells contain nuclei and some also contain abundant cyto- 



J show cytoplasmic inclusions, but these tissues pro- 

<iuce a negative Feulgen reaction; evidently the cellular contents 

American Fern Journal 

Volume 60, Plate 11 



Tracing of a Composite Photograph of the Vascular Cylinder of 
Lycopodilt^i fl.\belliforme. For abbreviations see Plalr 9. 

Anatomy of Lycopodium 


































c3 o3 

s s 

«1 w 

^ =1 

O O 


3. 3. 




a. a. 

















■S* ^ 

c c 

5 = s 

S c 







O 1= 









n " 





t ^^ 














































h3 h4 















American Fern Journal 

Volume 60, Plate 12 

fj PAR 



100 u 

Tracings of Composite Photographs of the Vascular Cylinders of 
Lycopodium clavatum and L. tristachyum. For abbreviations see Plate 9. 

Anatomy of Lycopodium 97 

are not nuclear material. 

This investigation shows that in addition to the basic stelar 
patterns known to exist in the genus Lycopodium, as described 
by Holloway (1919), there is a difference in size, shape and orienta- 
tion of the xylem and phloem tissue in the six species herein 
studied. The differences are characteristic of each species, and in 
anatomical studies they could be used as criteria for species 

Literature Cited 

Bower, F. O. 1935. Primitive land plants. Macmillan, London. 
Cardillo, F. M. 1966. The comparative anatomy of six north-eastern species 

of Lycopodium. Ph.D. Dissertation, St. Bonaventure Univ., St. 

Bonaventure, N. Y. 
. 1967. The ecology and distribution of the genus Lycopodium in 

southern Cattaraugus County, New York. Science Studies 24:35^3. 
Holloway, J. E. 1916. Studies in the New Zealand species of the genus 

Lycopodium. Part I. Trans. New Zealand Inst. 48:253-303. 
— . 1919. Studies in the New Zealand species of the genus Lyco- 

podium. Part IIL Plasticity of the species. Trans. New Zealand 

Inst. 51:161-216. 
Jensen, W. A. 1962. Botanical histochemistry. W. H, Freeman, San Francisco, 
Roberts, E. A. and S. D. Herty. 1934. Lycopodium complanatum var. 

flabellifornie Fernald: its anatomy and methods of vegetative propa- 
gation. Amer. J. Bot. 21:688-697. 
Wardl.\w, C. W. 1965. Organization and evolution in plants. Longmans^ 

Green, London. 
WiLCE, J. H. 1965. Section Complanata of the genus Lycopodium. Nova 

Hedwigia Beih. 19:1-233. 

Department of Biological Sciences, Quincy College^ 
QuiNcY, III. 62301. 

98 American Fern Journal 

A Note on the Gametophytes of Cystopteris tenuisecta 

S. S. BiR 

Taxonomically the genus Cystopteris Bernh. was divided into 
two subgenera, Cystopteris and Acystopteris, by Blasdell (1963). 
Gametophytes of several members of subg. Cystopteris^ such as C. 
douglasiij C. protrusa, C. diaphana, C, hulhijera^ and C. fragilis 
(Blasdell, 1963; Kaur, 1963; Bir & Trikha, 1970), have been 
studied, but up until now no observations have been made on 
the gametophytes of members of subg. Acystopteris, to which the 
Asiatic species C. tenuisecta (Blume) Mett. belongs. Furthermore, 
this species has also been referred to Cornopteris by Tardieu-Blot 
(1958). Therefore, the present study was made to find out the 
relationship of C. tenuisecta to the other members of the genus 
and also to see if it really belongs to Cystopteris or not, 

Cystopteris tenuisecta grows luxuriantly along open roadsides 
at Tiger Hill (alt. 2,400 m), Darjeeling (Eastern Himalayas). 
Both tetraploid sexual {n = 84) and triploid hybrid (2n = 126) 
individuals are abundant (Bir, 1961), The present study concerns 
only spores collected from tetraploid plants because the spores 
of the triploid did not germinate even after several trials. 
Gametophytes were raised by sowing the spores on full strength 
as well as on half-strength Knop's Solution. These were trans- 
ferred to steriUzed garden soil in pots when they were just 

Freshly collected spores take about five or six weeks to germi- 
nate. Early stages of germination are very like those described 
earher for other leptosporangiate ferns, such as Hypodematium 
crenatum (Loyal, 1960) and Diplazium (Athyrium) esculentum 
Olehra, 1949; Nayar, 1960). On germination the exine ruptures 
irregularly near the central region of the spore, and a green germ 
filament emersres. fnllowprl hv flip fir«f rKi^zr^irl fti^rvic^f^moc ^iie^v 

h 2). 


4-6-celled, an apical 

cell is cut ofif by formation of two oblique walls in the terminal 

American Fern Journal 

Volume 60, Pl.vte 13 

Fig*. 3-« 

Gametophytes of Cystopteris tenuisecta. Figs. 1-4. Filamentous 

Prothallus in CnOWD^JiCTTL- 

TURE. Fig. 7- M ARniiMAi. papillate hair. Fig. 8. Froth 

100 American Fern Journal 

cell {Fig. 3), In crowded cultures much elongated filaments are 
produced before an apical cell is organized {Fig. 4)- Lateral expan- 
sion of the prothallus takes place subsequently by longitudinal 
division in the penultimate cells of the germ filament and by suc- 
cessive lateral divisions of the apical cell on both of its sides. 
These divisions give rise to a spathulate prothallus. The apical 
cell is soon replaced by an apical meristem, and this results in 
formation of a typical, cordate prothallus (Fig. 5) with an apical 
notch, except in those cases where the cultures are crowded 
{Fig, 6). Usually the cordate condition is attained in about three 

The mature gametophytes resemble closely those of other higher 
leptosporangiate ferns. The central cushion is two or three cells 
thick. The margin is smooth. Marginal hairs start developing 
after the apical meristem is established and the prothallus becomes 
notched {Fig. 8), After this, superficial hairs develop generally on 
the prothallus surfaces. The hairs are unicellular, papillate, chloro- 
phyllous, and non-glandular {Fig. 7). The sex organs appear when 
the gametophytes are about four months old, and their develop- 
ment is of the usual type. 

The gametophytes of C. tenuiseda resemble those of C. fragilis 
(Kaur, 1963) and C. douglasii (Blasdell, 1963) in having a smooth 
margin and short hairs that appear late in gametophyte develop- 
ment. The gametophytes differ from those of C. diaphana, which 
have irregular margins at maturity and longer hairs that appear 
soon after the cell plate stage is initiated. The marginal, non- 
glaii3ular hairs on the prothallus of C. tenuiseda are similar to 
those on the prothalli of Diplaziopsis javanica (Bir, 1970), Atkyrium 
japmicum, A, thelypterioides (unpubl. data), and Hypodematium 
crenaium (Loyal, 1960). On the basis of gametophyte characters, 
C. tenuiseda is correctly placed in Cystopteris subg. Acystopteris, 
not in Cornoptens, Furthermore, Cystopteris is closely allied to the 
Athyrioid ferns that have close affinities to the Dryopteroid ferns. 

Literature Cited 

Bir, S. S. 1961. In P. N. Mehra, Chromosome mimbers in Himala^^an ferns. 

Res. Bull. Panjab Univ., n.s. Sci. 12:139-169. 



sis javauica (Bl.) C. Chr. Plant Sci. 1 :(in press). 
— . and C. K. Trikha. 1970. MorDholocical ol 

Cystopteris Bernhardi. Nat. Acad. Sci. India, (in press). 
Blasdell, R. F. 1963. A monographic studj- of the fern genns Cystopteris. 


Mem. Torrey Bot. Club 21(4) :1-102. 

Sci. 58:148-152. 

Loyal, D. S. 1960. Some observations on the gametophytes of Hypodematium 

crenatum (Forsk.) Kuhn with a note on the phyletic affinities of the 
genus. J. Indian Bot. Soc. 39:133-139. 

]Mehba, p. N. 1949. Observations on the prothallus of Diplazium esculent um. 

J. Indian Bot. Soc. 28:1-8. 
Nayar, B. K. 1960. The gametophyte and young sporophyte of Athyrium 

esculentum. Amer. Fern J. 50:194-203. 
Tardieu-Blot, M.-L. 1958. Validation of some new combinations. Amer. 

Fern J. 48:31-34. 

Department of Botaxy, Punjabi University^, Patiala, 


C. V. Morton 

Anogramma leptophylla (L.) Link (Fil. Sp. 137. 1S41) is a rather 
common and widespread fern occurring in both the Old and New 
Worlds. It is based on Polypodium leptophyUmn L. (Sp. PI 1092. 
1753). So far as I know, no one attempted to pick out a lectotype 
for this species until Pichi-Sermolli (Webbia 21: 497-502. 1966). 
However, I am unable to agree with the lectotype chosen or the 

method of selection. 
The original pubUcation is as follows: 

leptophyllum 46. Polypodium Pfrondibus bipinnatis: steriUbus brevissimis; 

pianis cuneiformibus lobatis. Magn. monsp. 5. 1, o. 
Adiantum minimum, foho vario. Magn. monsp. 5, t. 5. 
Adiantum filicinum leptophyllvmi elatius hispanicum. Ban. 

rar. 1210 t. 431, opt. . . 

Habitat in Ilispania, Lusitania, Galloprovmcia. 

Pichi-Sermolli wrote: "Lirni^ gives Magnol's and Barreher's 
phrase names and adds no original description of the species, as he 

102 American Fern Journal 

does for the new species such as Poly podium varium (p. 1090), 
Polypodium marginale (p. 1091), etc. Thus we are entitled to 
beUeve that Linne did not intend to introduce a new species, but 
accepted the basic concept of the species as established by Magnol 
and Barrelier. Consequently in selecting the lectotype, we have 
to take into consideration the plants described by these authors. 
However, it is worth mentioning that Linn6 notes that Barrelier's 
plate is 'opt.,' this means that he knew the species, probably 
through the specimen kept in the Linnean herbarium." Later he 
wrote: "In conclusion, Anogramma leptophylla cannot be typified 
on the specimen of the Linnean herbarium although present in it 
before 1753, since Linn^ did not describe the species at all, but 
clearly accepted the basic concept of the species as established by 
Magnol and Barrelier." 

These remarks show a basic misunderstanding of the process of 
lectotypification of Linnean species as established by generations 
of workers and brought together by Stearn in his preface to the 

Plantarum." Pichi-Sermolli 

only a few of the species described in the "Species Plantarum" 
were new and that the others are to be typified on the basis of the 
literature reference cited is contrary to the Code. Botanical no- 
menclature begins with 1753 and all of Linnaeus' species are con- 
sidered as new as of that date. Their lectotypification goes forward 
and depends on the work of later authors and does not go back- 
ward into pre-Linnaean literature necessarily. There is a general, 
if tacit, agreement that Linnaean species ought to be lectotypified 
on the specimens in the Linnean Herbarium whenever possible, 
that is, when there is reason to believe that a specimen was in the 
herbarium in 1753 and when it agrees with the diagnosis. 

In the case of Polypodium leptophyllum L. there is a specimen 
m the Linnean Herbarium in London, under the number 1251.56. 
It bears the name Polypodium heterophyllum in Linnaeus hand and 
the number 46. As Pichi-Sermolli remarks, it is clear that Linnaeus 
first called this P. heterophyllum and altered the name in the publi- 
cation to P. leptophyllum because he found that he had already 
applied the name P. heterophyllum to another soecies. The num- 

Taxonomic Notes on Ferns 103 

ber "46" shows that this is indeed the species described as P. 


Pichi-SermolU has chosen as lectotype Toumefort 5337 in the 
Paris herbarium. BarreUer's herbarium was destroyed by fire, and 
so no specimen by him can be lectotype. This leaves iVIagnol 
(1697), who indicated that he had been given his specimen by 
Tournefort. No Magnol herbarium is known, and therefore Pichi- 
Sermolli goes back to the Tournefort Herbarium, where the plant 
now know^n as Anogramma leptophylla is represented by sheet 
5337, which is therefore designated as lectotype. But there is 
really no proof that this specimen has anything to do with the 
species — it was not cited by Linnaeus, it was not seen by Linnaeus, 
and in fact Linnaeus did not mention Tournefort at all; and there 
is no certainty at all that the plant that Tournefort gave to Magnol 
w^as really the same as this sheet in the Tournefort Herbarium. 
Therefore, Tournefort 5337 as a lectotype is impossible, especially 
Avhen there is a perfectly good specimen in the Linnean Herbarium 
to be the obvious lectotype. Therefore, I propose the sheet 1251.56 
in the Linnean Herbarium as lectotype of Polypodium leptophyllum 
L. and reject Tournefort 5337. 

National Museum of Natural History, Washington, D.C. 

Taxonomic Notes on Ferns, IV 

C. V. IMorton 

In the previous numbers of this series of casual notes on ferns, I 
published on Athyrium lilloi (Amer. Fern J. 50: 275, 276. 1960), 
on Thelypteris sect. Glaphyropteris, Blechnum penna-manna, and 
Asplenium varians (Amer. Fern J. 51: 37-39. 1961), and on 
Anopteris hexagona and Elaphoglossum denuclatum (Amer. Fern J. 

iQfio^ TViP rirpcpnt naner discusses four species of 



LoiL^RiA MicROPHYLL.^ Goldm. Nov. Act. Acad. Caes. Leop. 

CaroL 16, Suppl. 1: 460. 1843 = BlecJinum microphyllum 
(Goldm.) Morton, comb. nov. 

104 American Fern Journal 


austr. Aiidibus ad Antuco, Lect. 1828/' Poeppig, The holotype in the 


doubted isotype in the herbarium in Berlin-Dahlem, marked in Kunze's 


photogi-aph 10182). This photograph shows that L. australis is not a 


"Index Filicum," but is the same as the species later described as Lomaria 

germatmt Hook. The epithet ''australis,'' although the oldest for this 
species, may not be used in Blechnum because of the existence of B, 
ausirale L. (1767). 

Lomaria gayana R^my & F6e in Gay, Hist. Fis. PoL Chile Bot. 6: 481. 
1853. Type: "Cordilleras de Talcaregue, provincia de Colchagua/' Chile, 

Blechnum poeppigianum Sturm, Enum. PL Crypt. Vase. Chil. 26. 1859. 
Based on Lomaria australis Kunze, non Blechnum ausirale L. 

Blechnum gayanum (R6my & F6e) Sturm, Enum. PI. Crypt. Vase. Chil. 
24. 1859. 

Lomaria germainii Hook. Sp. Fil. 3: 32, pi 152, 1860. Syntypes: Cerro de 
la Campana de Quillota, Germain and Antuco, Chile, Poeppig. Looser 
chose the Germain collection as lectotjrpe. 

Blechnum gerniainn (Hook.) Christ, Bull. Herb. Boiss. H, 2: 554. 1902. 



collected in Chile by Aleyen and marked ''Blechnum {Lomaria) 
microphyllum n. sp." (Morton photograph 10180). This is surely 
an isotype and possibly the holotype. I do not know the location of 



(Poir.) Kuhn, but is identical ^nth Blechnum gayanum (R^my & 
F^e) Sturm, as suspected by Gualterio Looser in his revision of 
the genus Blechnum in Chilei. However, since he had not seen the 
type, Looser placed Lomaria microphijlla Goldm. among the 
dubious species. In the "Index FiUcum," Lomaria gayana is also 
referred erroneously to B. penna^marina, but, as shown by Looser, 
it also belongs with the species that has been generally known as 

l—fe'^Mr ^"''^^'■s^**"^ (Universidad Catol. de Chile) 32(2): 7-105, M 

Taxonomic Notes on Ferns lOo 


Lomaria (jermatnit Hook, or Blechnum (jermainii (Hook.) Christ. 
Additional synonyms, according to Looser, are Blechnum andicola 
(Phil.) C. Chr. and B. araucana (Phil.) C. Chr. A Chilean specimen 
collected by Werdermann has been distributed under the unpub- 
lished name ^^ Blechnum nivale Reimers, u. sp. 

LoAiARiA SETiGERA Gaud, Ami. Sci. Nat. [Paris] 5: 9S. 1S23 
Blechnum magellanicum Desv. var. angustiseta (Bory) Mor- 
ton, comb, nov, 

Lomaria rnageUanica angusiiseta Bory ex Dumont d'Urville, Ann. Linn. 

Sue. [Paris] 4: 597. lS2o [1S26]. Based on Lonuxria sciigera Gaud. 
Blechnum magellanicum (Desv.) Mett. var. setigerum (Gaud.) C. Chr. Ark. 

for Bet. 10(2): 9. 1910. Based on Lomaria setigera Gaud. Incorrect, 

since the earliest varietal name was not adopted. 
Blechnum tahulare (Thunb.) Kuhn var. setigerum (Gaud.) Capurro, Anais 

Prim. Reun. Sul-Amer. Bot. 2 : 124. 1940. Incorrect, since the 

earliest varietal name was not adopted. 

Type: lies Malouines, i.e. Falkland Islands, Antarctic South 
America, Gaudichaud (holotype P, Morton photograph 4406). 

Ptvchopyllum magellanicum Klotzsch, Liuuaea 18: 533. 1844 
[May-Aug., 1845] = Hymenophyllum seselifollum K. B. Presl. 

Hynimophyllum magellanicum (Klotzsch) Kunze, Bot. Zeit. 5: 226. 184<. 
(wrongly attributed to "Willdenow")- 

It is unfortunate that the species well known in southern South 
America as H y me noph ijllum magellanicum (Klotzsch) Willd. 
must now be called H. seselifolium K. B. Presl, a name perhaps 
never used since its original publication. Presl published this 
species in his little book *'Hymenophyllaceae" which is dated 
"1843" but which probably came out in 1844 (see William T. 
Steam, J. Soc. Bibl. Nat. Hist. 3: 15. 1954). Klotzsch's name 
Ptychophyllum magellanicum is in a volume of Linnaea dated 
"1844," but Stafleu's recent and invaluable "Taxonomic Litera- 
ture" establishes the actual date of pubUcation as between May 
and August, 1845. Thus Presl's name has clear priority of a year 
or more. Klotzsch based his new species on a plant from the Straits 
of IMagellan in the Willdenow Herbarium (no. 20245) in Berhn 
and on a specimen from Chilo6 Island collected by PhilippL 
Presl's type came from Chilo^ also, a collection of Hugh Cummg 

106 American Fern Journal 

(no. 6, presumably in the Presl Herbarium in Prague). 


Desv. (]\Iem. Soc. Linn. Paris 6: 331. 1827), which was based on a 
collection merely indicated as from South America. The holotype 
is in the Desvaux Herbarium in Paris (Morton photograph 4596); 
it was very likely collected by Commerson. Desvaux and Klotzsch 
independently adopted the epithet ''magellanicum,'' a naturally 
appropriate name considering the locality involved. Although 
Desvaux' name is the earliest for this species, it may not now be 
transferred to Hymenophyllum because there is already the name 
Hymenophyllum magellanicum (Klotzsch) Kunze, which although 
taxonomically the same is not nomenclaturally the same since it 
is based on a different type. 

Mertensia magellanica Desv. Mem. Soc. Linn. Paris 6: 20L 


Desvaux attributed this name to Poiret, referring to Poiret in 
Lamarck. Encych Meth. SuppL 3: 669. 1813; however, at this 
place there is no species called Mertensia mageUanica. The epithet 
viagellanim must be derived from the Straits of Magellan, and 
the only species on the page cited by Desvaux that is from this 
locality is Mertensia quadripartita Poir. This is however only an 
inference, although a logical one, and so the name M, magellanica 
Desv. must be considered a nomen nudum, since there is no de- 
scription and no complete reference to a description under another 
name. That the inference mentioned above is correct is shown by 
a specimen in the herbarium of the Museum National d'Histoire 
Naturelle, Paris, where a specimen from the Desvaux Herbarium 


Mertensia ma 


quadripartitum Poir. (in Lam. Encycl. M^h. 5: 543. 1804), which 
was later called Mertensia quadripartita (Poir.) Poir. The name 
''M. magellanica'' was very likely merely a slip of memory on the 
part of Desvaux, but it has gotten into the literature. 

National :Museum of Natural History, Washington, D.C. 

^Morphology of Hypodematium 107 

A Reinvestigation of the Morphology 
of Hypodematium crenatum 

B. K. Nayar and Nisha Bajpai 

Taxonomically as well as phylogenetically the small fern genus 
Hypodematium remains a problematic one. It includes the wide- 
spread, pantropic species H. crenatum (Forsk.) Kuhn and perhaps 
three or more other species, each of limited range in the China- 
Japan region; some pteridologists detect as many as 16 species. 
The genus has been associated with the woodsioid, tectarioid, 
athyrioidj and thelypteroid ferns by different taxonomists, and in 
some cases even by the same taxonomist at different times. 
Morphologically the genus is little known except for some aspects 
of H. crenatum published by Hayata (1927), Mehra and Loyal 
(1956), Loyal (1960), and Iwatsuki (1964a, b). Since the descrip- 
tions of these authors contain some discrepancies, we thought 
it worthwhile to reinvestigate the morphology of H, crenatum and 
to see whether the morphology of this plant could yield some 
additional clues as to the relationships of the genus. 

Material for this study was collected growng in rock crevices 
from Chakrata (Central Himalaya) and was then cultivated in 
the fernery of the National Botanic Gardens. Spore morphology 
was studied using standard palynological techniques, as described 
by Nayar and Devi (1964). Prothalli were grown on steriHzed 
nutrient agar medium in petri dishes maintained at 24 ± 2° C 
and at about 600 ft-c light intensity, as described by Nayar (1962). 
The hght source was fluorescent ''day light'* lamps for 12 hours in 
each day. Vascular organization of the rhizome was studied 
from steles excavated mechanically from the rhizome, as well as 
from reconstructions ba.sed on camera lucida drawings of serial 

microtome sections. 

The rhizome is thick, creeping, cylindrical, bears several 
dormant branches on the ventral surface, and is densely covered 



late, glandular hairs {Fig. 7). The apex of the palea is uniseriate and 

American Fern Jourxal 

Volume 60, Plate 14 

Morphology of the Sporophyte of IIypodematium crenatum. Fig. 1- 

5AL VIEW. Fig. 2. 

Vascular cylinder of a portion of the rhizome. 
Same, lateral view. Fig. 3. Mature palea. Fig. 4. Same, apex. Fig. 5. 
Same, margin. Fig. 6. Immature palea at stage marginal hairs are 
produced. Fig. 7. Papillate hairs on rhizome. Fig. 8. Lower leaf epi- 
dermis. Fig. 9. Upper leaf epidermis. Fig. 10. Portion of indusium 
showing acicular hairs. Fig. 11. Mature sporangium. Fig. 12. Immature 


m SHOWING origin OF THIRD ROW OF STALK CELLS. The abbrevia- 
tions are: a = acicular hair, b = glandular hair, e = secondary row 






is terminated by a non-glandular, barrel-shaped cell {Fig. 4)- 
Young paleae bear sparse, unicellular^ papillate, glandular hairs 
{Fig, 5) on the margin and both surfaces. The hairs are formed 
early during development of the paleae, sometimes when the paleae 
are only three cells broad {Fig. 6), and are deciduous. Filamentous, 
marginal protrusions bearing hairs are found occasionally on the 
basal region of the paleae. The rhizome is parenchymatous, with 
small, globose or ovoid masses of thick-walled cells with dark 
brown contents scattered irregularly in the ground tissue. In 
rhizome transections these sclerenchyma masses appear Hke the 
sclerenchyma strands found in many dryopteroid ferns, but they 
are not elongate strands. The vascular cyhnder of the rhizome 
{Figs, 1, 2) [^ distinctive and, as reported by Alehra and Loyal 
(1956) and Iwatsuki (1964a, b), has a broad, undivided ventral 
half and a dissected dorsal half, recalling that of the Lomariopsi- 
daceae (Xayar & Kaur, 1965). In the adult rhizome there are 
generally three irregular rows of leaf gaps dissecting the dorsal 
half of the stelar cylinder— not two alternating rows, as reported 
by Iwatsuki (1964a, b). The leaf gaps are clearly arranged in a 
spiral around the vascular cylinder, 's\nth a larger distance between 
successive gaps on the ventral side. The lateral leaf gaps are 
markedly displaced towards the dorsal median plane. The leaf 
gaps are large and spindle-shaped and dissect the dorsal half of 
the stele into narrow, ribbon-like vascular strands. Leaf traces 
{Figs, 1, 2, i) are paired, ribbon -like strands, and the leaf gap 
extends conspicuously on the abaxial as well as on the adaxial 
side of them. Root traces are restricted to the ventral, undissected 
portion of the stele, and originate as thick, superficial, cylindrical 
branches; they are not associated with the leaf gaps. Branch trac^es 
{Fig. 2, k) are profuse and are restricted to the ventral half of the 
stelar cylinder. They are similar to the root traces in being cylin- 
drical, solid strands arising superficially on the stelar surface (not 
associated with any branch gap) and oriented in the same way 
(not at the margins of the ventral meristele, as :Mehra and Loyal 
(1956) reported). While passing through the cortex of the rhizome, 
the branch trace becomes meduUated and at the branch base it is 

110 Amekican Fern Journal 

siphonostelic. In developed branches it becomes dissected into a 

with soirallv arran 

but as the branch thickens and grows away from the main rhizome, 
the ventral leaf gaps become displaced and the ventral half of 
the vascular cylinder becomes undissected. The xylem tissue of the 
vascular cylinder is massive and often has 10-12 rows of tracheids 


The phloem forms a thin sheath around the xylem, but is discon- 
tinuous at the region of the leaf gaps (margins of the ribbon-like 
meristeles). The pericycle is three- or four-layered, and the endo- 
dermal cells occasionally have faintly thickened inner walls. 

The leaves are crowded, with the leaf bases persistent and 
densely aggregated on the dorsal surface of the rhizome. The 
leaf bases are conspicuously swollen and tuberous, as reported in 
Pteris wallichiana (Chandra & Nayar, 1970). ]Many of the lateral 
leaves are undeveloped except for their swollen bases; the upper 
part of such leaves is represented by a deciduous, hump-like pro- 
trusion. The swollen leaf bases bear profuse paleae and unicellular 
hairs. They are parenchymatous hke the rhizome and possess scat- 
tered sclerenchyma masses; dense deposits of starch occur in the 
parenchyma cells. The stipe above the swollen base is sharply 
delimited, nearly cylindrical, and hard due to a broad hypodermal 
sheath of 5-8 layers of very thick-walled cells. The epidermis 
is thin-walled and bears unicellular hairs. Sclerenchyma masses 
are absent in the stipe. The vascular supply consists of a pair 
of broad, laterally placed strands, with the inner walls of the endo- 
dermal cells conspicuously thickened and dark brown. The leaves 
are shed where the stipe joins the swollen leaf base, but, as noted 
by Iwatsuki (1964a, b), there is no clear abscission tissue such as 
occurs in ferns ^vith distinctly articulated leaves (e.g., the Poly- 
podiaceae). A layer of cells of the stipe next to the swollen base 
turns brown in the old leaves and ultimately shrivels up, resulting 
in the shedding of the anterior part of the leaf. 

The rachis and its branches are subterete \dxh a more or less 
flat adaxial surface. The two vascular strands of the stipe fuse by 
their abaxial margins into a solitary bundle, which is nearly 

Morphology of Hypodematiuai 111 

trigonal in the anterior half of the rachis. Xylem tissue of the rachis, 
bundle is deeply adaxially channeled (U-shaped), with broad, 
thin, prominently incurved, flap-like margins. Phloem entirely 
surrounds the xylem in a thin sheath, and the pericycle which 
surrounds the phloem is one cell thick, except on the adaxial 

side, where it is much thicker and fills the concavity of the vascular 


The lamina is herbaceous and decompound. The ultimate 
segments are oblong, with blunt apices and a broadly lobed margin. 
Venation is pinnate, with pinnately branched main lateral veins 
corresponding to the marginal lobes. Elongate, unicellular or 
occasionally bi- or tri-cellular, acicular, achlorophyllous, thick- 
walled hairs, often with peg-like protuberances on the inner surface 
of the cell Avail (Fig, 8), are found all over the lamina, more 
profusely on the veins and axes. Epidermal cells of the lamina, 
other than those over the veins, are thin-walled, chlorophyllous, 
and ANith a broadly sinuous contour; cells of the lower epidermis 
have a more conspicuously sinuous outhne than those of the 
upper epidermis. Epidermal cells over veins (Figs. 8, 9, m) 
on both surfaces differ in being achlorophyllous, narrow with 
a smooth contour, and conspicuously elongate parallel to the 
vein. Similarly, the cells at the margin of the lamina {Fig. 9, 
w) are achlorophyllous, with smooth contour, thickened peripheral 
walls, and are elongate parallel to the margin. This band of 
specialized marginal cells is continuous along the margin of 
the pinnules and becomes more than one cell broad at the basal 
region of the pinnules and at the sinuses between the marginal 
lobes, recalling the sinus-membrane of the thelypteroid ferns. 
The bands of elongate, achlorophyllous epidermal cells above 
the veins extend to the margin and merge with the marginal band. 

The sporangia are produced in circular sori borne medianly on 
the lateral veinlets. The sorus is protected by a deeply reniform, 



those on the lamina, but often having hooked apices, are profuse 
on the outer surface as well as on the margin (Fig, 10) ; the marginal 

American Fern Journal 

AoLUME 60, Plate 15 

Morphology of the Gametophvte of Hypodematium cbenatum. 
Figs. 13-19. Stages in prothallial plate development. Fig. 20. Apex of 

YOUNG cordate PROTH- 

IS. Fig. 21. Adult prothallus. Fig. 22. M 



Fig. 25. Antheridium. Fig. 26. Adult prothallus, anterior margin with 
HAIRS. The abbreviations are: a == 


and q = SPOHE coat. 


Morphology of 113 

hairs often are larger than the superficial ones. Sporangia {Fig, 11) 
are the typical leptosporangiate type, with a long stalk composed 
of four tiers of elongate cells. The stalk is two cells thick, with the 
rows contiguous with either end of the annular ring, except at the 
capsule base, where there is a third row two cells long formed 
secondarily during sporangial development as a downward out- 
growth of the basal cell of the capsule wall on one of its flat sides 
{Fig, 12). The annulus consists of 16-20 indurated cells. 

The spores are monolete and bilateral with a smooth, dark 
brown exine and loose, densely granulose perine that is wrinkled 
into short, thin, sparse, lobate folds. The spores average 32 ^ 
X 45 )it (P X E, exclusive of perine) and are planoconvex in 
lateral view and oblong in polar view. The laesura is about 30 /a 
long and crassimarginate. Spore germination is the Vittaria-type 
(Nayar & Kaur, 1968). 

The prothallia develop basically like the Aspidium-tyY^e 
(Nayar & Kaur, 1969), and exhibit a full range of variation with 
respect to early formation of hairs. In the majority of cases the 
terminal cell of the germ filament becomes quiescent and produces 
a crowning unicellular hair {Figs, 13, 14). A lopsided prothaUial 
plate is developed by the activity of the penultimate cells {Figs. 
15-17). A second hair is developed on the more expanded side of 
the plate, and this is usually followed by the estabhshment of a 
lateral meristematic cell between the two hairs. The gro\nng 
apex of the prothallus soon becomes symmetrical, with a median 
meristematic cell {Fig. 18). Many thalli omit a meristematic cell 
stage. In a few cases hair formation is rather delayed during pro- 
thaUial development, and a prothalhal plate is formed by the ac- 
tivity of the terminal cell of the germ filament {Figs. 15Aj 16A). 
In such cases the young prothallial plate is symmetrical and 
naked. Soon, however, a terminal hair develops on the plate and 
the plate becomes asymmetric due to unilateral growth. The 
prothalli become distinctly cordate about 40 days after spore 
germination, A pluricellular meristem replaces the meristematic 
cell (or in thalli which skip the meristematic cell stage, a meristem 
is established directly from anterior marginal cells) when the apex 

114 American Fern Journal 


of the thallus becomes cordate (Figs. 19 , 20). Young thalli are 
profusely hairy. The hair-bearing marginal cells frequently become 
protruded and sometimes form short protuberances two cells 
long, making the prothallial margin irregular {Fig, 20), 

The prothalli reach maturity about 60 days after spore germina- 
tion. The adult prothallus {Fig, 21) is of the common, cordate- 
thalloid type; it is symmetrical^ nearly circular, 6-7 mm across, 
and has a median midrib 6-8 cells thick, and broad, flat wings. 
The wing cells possess collenchyma-like thickenings at the corners 
{Fig, 24)- Unicellular papillate hairs are profuse all over the thallus; 
the hair-bearing marginal cells are prominently protruded. The 
hairs are thin-walled, chlorophyllous (with chloroplasts markedly 
smaller than those in other prothallial cells), and secretory, the 
secretion forming a prominent, tough, w^axy cap crowning the hair 
{Fig, 22) . In addition, all the adult prothalli (not some of them only, 
as Loyal (1960) reported) bear profuse, much elongate, rhizoid- 
like hairs (''bristles") on the anterior margin and surfaces. These 
are thin-walled, unicellular {Figs, 23, 26), and sparsely chlorophyl- 
lous Uke the papillate hairs, but are several times longer, more 
highly vacuolated, and devoid of any secretion. Occasionally both 
types of hairs have one or two transverse septae in their basal 
half. Hairs intermediate in size between papillate hairs and 
rhizoid-like hairs also occur, and these commonly have a scanty 
secretion. ''Acicular hairs similar to those on the lamina of fronds" 
that Loyal (1960) reported were not found on the prothallus. 

Sex organs are of the common type found in advanced lepto- 
sporangiate ferns. The antheridium is subglobose with a funnel- 
shaped basal cell {Fig, 25). The cap cell collapses at antheridial 



reported. The neck canal cell is binucleate and swollen toward 






jMorphology of Hypode.matium 115 

the thelypteroid ferns by Ching (1933, 1935, 1940, 1963), the 
athyrioid ferns by Loyal (I960), and the tectarioid ferns by 
Holttum (1954) and others. The acicular foliar hairs and paired 
leaf traces are construed as indicative of thelypteroid affinity, 
the general morphology of the lamina and its resemblance to 
Lastreopsis as indicative of tectarioid affinity, and the paired 
leaf traces and basic chromosome count {n = 41) as indicative of 
athyrioid affinity. The chromosome count alone seems sufficient 
to rule out a thelypteroid relationship, despite the acicular foliar 
hairs of the thelypteroid type on H, crenatum. Hypodematium 
includes species which lack acicular hairs, and so this feature 
should not be given much weight in phyletic considerations, as 
Iwatsuki (1964a, b) pointed out. Also, characteristically the paleae 
of thelypteroid ferns also bear acicular hairs, which are absent 
in H, crenatum; the paleae of which bear instead papillate hairs 
both on the margin as well as on the surface, as in some of the 
dryopteroid and athyrioid ferns. The prothalli of thelypteroid 
ferns have DrynariaAype development, in contrast to the Aspi- 
dium-type characteristic of Hypodematium and the dryopteroid 
and tectarioid ferns. At the same time, Hypodematium cannot be 
held as closely related to the latter groups, mainly because of its 
binary leaf trace with broad, ribbon-like vascular strands. Both 
the dryopteroid and tectarioid groups have a large number of 
slender, cylindrical vascular strands constituting their leaf trace. 
In addition, the prothaUial trichomes, particularly the unique, 
elongate, rhizoid-like, non-glandular hairs of Hypodematium^ have 
no parallel in any of these ferns. The tectarioid ferns, as far as 
known, possess a few branched, club-shaped glandular hairs on 
their prothalli, in addition to papillate secretory hairs. The 
woodsioid as well as the dryopteroid ferns have only papillate 
hairs. Hypodematium resembles the athyrioid ferns in having paired 
leaf traces and two ribbon-hke vascular strands in the stipe, 
but differs in its characteristic stelar architecture of the rhizome 
and more significantly in its prothaUial morphology; the athyrioid 
ferns have a Drynaria-type, not an Aspidium-iype, of development. 

Hypodematium^ therefore, cannot be held to belong to any of the 

IIG American Fern Journal 

groups of ferns with which it has been associated in the past by 
different pteridologists. However, there is little reason to doubt 
that it is most closely allied to the Dryopteridaceae, as is amply 
proved by spore and prothallus morphology and chromosome 
number. That it is an evolutionarily advanced genus is evident 
from its stelar morphology. It seems clear that the stelar cylinder 
with leaf gaps restricted to the dorsal surface is derived from a 
typical dictyostelic condition by displacement of the ventral 
leaf gaps in response to the peculiar habit of the plant (growing 
wedged in crevices of hard rock and firmly attached to the sub- 
stratum), and is not a primitive condition, as suggested by Iwatsuki 
(1964b). That the ancestral condition is a radially symmetrical 
dictyostele is indicated by the presence of such a vascular cylinder 
at the bases of rhizome branches, which gradually progresses to 
the fully developed form by displacement of ventral leaf gaps. 
The presence of suppressed leaves on the rhizome is another such 
indication. It is only the lateral leaves towards the ventral surface 
that are suppressed, and not any of the median ones. Occasional 
suppression of some of the ventral leaves and displacement of 
ventral leaf gaps towards the dorsal surface occur in some genera of 
the Dryopteridaceae which include some species with erect 
rhizomes and others with creeping or semi-erect rhizomes, like 
Polystichum (Chandra, 1969) and Tedaria, A similar suppression 
of ventral leaves is encountered also in the creeping species of 
Oleandra (Nayar et al., 19GS). Also, the leaf gaps of Hypodemalium 
exhibit a distinct spiral arrangement on the stelar cylinder; they 
are not in tw^o or three dorsal, longitudinal rows, as described by 
jMehra and Loyal (1956) and Iwatsuki (1964a, b). A unique feature 
of Hypodemalium is its branch traces, which are solid, superficial 
strands on the ventral surface of the rhizome and resemble root 
traces. Such branch traces are not reported in any other fern with 
w^hich Hypodematium has been associated. They recall the stolons 
produced by Nephrolepis. 

It seems best to regard Hypodematium as an isolated genus in 
the Diyopteridaceae, as suggested by Nayar (1970). It represents 
a line of evolution coordinate with the drvooteroid, athyrioid, 

Morphology of hypode:matium 117 

and tectarioid ferns, but distinct from them. The latter three 
groups and Hypodemafmm each may be considered a subfamily of 
the Dryopteridaceae. 

Literature Cited 


ceae: Polystichoid and Thelypteroid Fems. Ph.D. Thesis, Agra Univ. 
Chandra, S. and B. K. Nayar. 1970. Some aspects of the morphology of the 

rhizome of Pteris wallichiana. Proc. Indian Acad. Sci. 71:79-85. 
Ching, R. C, 1933. On the nomenclature and systematic position of P0I3'- 

podium dryopteris L. and related :ipecies. Contr. Biol. Lab. Sci. Soc. 
China 9 :30-40. 

— ^ . 1935. On the genus Hypodematium Kunze. Sunyatsenia 3:3-14. 


senia 5:201-268. 

— . 1963. A reclassification of the family Thelypteridaceae from the 

mainland of Asia. Acta Phytotax. Sinica 8:289-335^ 

CoPELAND, E. B. 1947. Genera Filicum. Chronica Botanica, Waltham, Mass. 

Hayata, B. 1927. On the systematic importance of the stelar system in the 

Filicales, Bot. Mag. Tokyo 41:697-718. 

HoLTTUM, R. E. 1954. Flora of Malaya, vol. 2. Ferns of Malaya. Gov't. 

Printer, Singapore. 

IwATSUKi, K. 1964a. On Hypodematium Kimze. Acta Phytotax. Gebot. 

21 :43-54. 

■ — . 1964b. Taxonomy of the thelypteridaceous ferns, with special 

reference to the species of Japan and adjacent regions. IL Circum- 
scription of the group. Mem. Coll. Sci. Univ. Kyoto, B, 31 :1-10. 

Loyal, D. S. 1960. Some observations on the gametophyte of Hypodematium 

crenatum (Forsk.) Kuhn with a note on the phyletic affinities of the 
genus. J. Indian Bot. Soc. 39:133-139. 

Mehra, p. N. and D. S. Loyal. 1956. Some observations on the cytologj- 

and anatomy of Hypodematium crenatum. Curr. Sci. 25:363-364. 

Nayar, B. K. 1962. :\Iorphology of the spores and prothalli of some species of 

the Polypodiaceae. Bot. Gaz. 123:223-232. 

— . 1970. Classification of the homosporous ferns. Chapter II in 
B. K. Nayar and S. Kaur, Companion to Beddome's Handbook to 
the Ferns of British India, Ceylon & Malay Peninsula. Chronica 

Botanica, Delhi. 

— , and S. Devi. 1964. Spore morpholog}- of Indian fems. I. Aspidia- 

ceae. Grana Palynol. 5:80-120. 


Egenolfia. I. The adult sporophytes of some species. J. Linn. Soc. 
Bot. 59:127-140. 

118 American Fern Journal 

— , and . 1968. Spore germination in homosporoiis ferns. 

J. Palynol. 4:1-14. 

— , and . 1969. Prothallial development in homosporous 

ferns, Phytomorphology 19:125-135. 

— . N. Bajpai, and S. Chandra, 1968. Contributions to the mor- 

phology of the fern genus Oleandra. J. Linn. Soc. Bot. 60:265-282. 

National Botanic Gardens, Lucknow, India. 

Shorter Note 

Pyrrosia princeps, a Fern New to Cultivation. — I have 
recently received for identification from Mr. J. W. Peterson an 
unidentified fern that is being cultivated in Longwood Gardens 
from material collected by R. G. Wilson in New Guinea and 
Bougainville (Peterson J -1087, collected Jan. 12, 1970). It proves 
to represent a species previously unknown in cultivation and very 
restricted in its natural distribution; 
Pyrrosia princeps (Mett.) Morton, comb. nov. 

Polypodium princeps Alett. Ann. Lugd. Bat. 2: 232. 1866. Type: New 

Guinea, Zi'ppe/ (holotype L, Morton photograph 833). 
Cyclophorus princeps (Mett.) C. Chr. Ind. Fil. 200. 1905. 

This is a characteristic species, known to me from only three 
collections (aside from the type) from New Guinea: Auga River, 
in clumps among rocks near river, alt. 580 m., Papua, Sept.-Nov., 
1933, Brass 5492; terrestrial in edge of forest on riverbanks, 
Biniguni Camp, Gwariu River, ISlilne Bay District, Papua, July 
30, 1953, Brass 23703; on limestone cHffs, Biak Island, December 
2, 1945, Grether & Wagner 4220, From the data on the label of the 
cultivated plant it appears that the species may occur also on 
Bougainville Island, Solomon Islands. 

In habit this species is quite different from the commonly 
cultivated Pyrrosia lingua (Thunb.) Farwell, which has long- 
stalked fronds scattered along an elongate, creeping rhizome. The 
rhizome is compact and larger in P. princeps j and the fronds are 
fasciculate. The fronds are probably the largest in the genus 
Pyrrosia, reaching more than a meter in length and 10 cm. in 

Recent Fern Literature 119 

width. The fronds are essentially exstipitate, which distinguishes 
them from such species as P. grandissima (Hayata) Ching. The 
Indian Pyrrosia costata (K. B. Presl) Iwatsuki and the Philippine 
P. splendens (K. B. Presl) Ching are similar in the frond outline, 
but the indument is quite different, the scales being dimorphous, 
the layer of peltate scales being underlain by white, dissected 
scales with elongate frizzy arms. In P. prmceps the scales of the 
lower surface are monomorphous, all like, with numerous, short 
spreading arms,— C. V. Morton, National Museum of Natural 
History, Washmgton, D.C, 20560. 

Recent Fern Literature 

A Commentary on Some Type Specimens of Ferxs in the 
Herbarium of K. B. Presl, by R. E. Holttum. Nov. Bot. Inst. 
Univ. Carol. Prag. 1968 :3-57. June, 1969.— The study of fern types 



certain extent neglected until the twentieth century. The earliest 
paper that I recall specifically on fern types is that of Christensen 
in 1910 on some of the types of Swartz. Probably no others were 
published until 1936, when Weatherby wrote a most valuable paper 
on the American types of Desvaux, and 1937, when Christensen 
wrote on the types of Cavanilles and Cesati. In 1954 I began a 
general study of fern types, which has continued up to the present 



The Czech botanist K. B. Presl described many species between 


have almost never been consulted, which is unfortunate since 
some of his species, although rather fully described for his day, 
have remained dubious. Dr. Holttum has remedied this by jour- 
neying to Prague to locate the INIalaysian types, which are among 
the most important, for most of Presl 's new species were based 
on the Philippine Island collections of Thaddeus Haenke and 
Hugh Cuming. Presl's herbarium was still unmounted and just 
as Presl left it when Underwood visited it in 1905, but it is now 
mounted and filed, partly in the general herbarium of the Bo- 

120 American Fern Journal 

tanical Institute of Charles University and partly in the National 
Museum, Prague. Holttum gives an interesting account of the 
history of the herbarium and also of the Haenke collections 
obtained on the Malaspina Expedition (1789-1794). Holttum's 
paper gives a commentary on all of PresPs Malaysian collections 
and gives his current disposition of them. The paper contains a 
rather large number of typographical errors, which are not at all 
Holttum's fault but which are attributable to the Czech type- 
setters' lack of familiarity w4th English and with some standard 
botanical practices, such as the setting of new combinations in 
boldface. The most significant errors I have noted are: 
Page 3, paragraph 1, lines 11 and 12 are reversed. 
Page 21, line 16: For '^Now identified as Drynaria quercifolia 
(Linn.) J. Sm.", read ''Now identified as Goniopteris tetragona 
(Swartz) Presl." I obtained the correct reading from Holttum 
himself, who consulted his original manuscript. 
Page 36, line 12: For ^1962," read '1862." 
The points of interest are far too numerous to mention here, 
but special mention should be made of the discussion of Presl's 
genera Pronephrium and Proferea, the former being an earlier 
name for Abacopteris F6e, if that is recognized as a genus, and the 
latter a synonym of Cyclosorus megaphyllus (Mett.) Ching 
(= Thelypteris Tnegaphylla (Mett.) K. hvatsuki). Holttum's in- 
tensive study of the ferns of this area for more than 40 years 
renders him uniquely qualified to discuss the taxonomy. I w^ould 
like to comment only on the treatment of five species, which are 
mentioned below* 

Poly pod 

alter nifolia (W 

todes nigrescent (Blume) J. Smith) . This misidentification of 
Polypodium allernifolium Willd. wath Polypodium nigrescens Blume 
is a mistake of Copeland. Sledge (Bull. Brit. Mus. Nat, Hist., 
Bot, 2 : 144. 1960) has commented that a photograph of the type 
in the Willdenow Herbarium in Berlin shows that alternifolium 
Willd. is a form of Polypodium phymatodes (i.e. P. scolopendria) 
rather than a synonym of P. nigrescens. He says that the type 

Recent Fern Literature 121 

sheet is a small but fertile frond, by which he must have been 

Willd. 19637/1 


19637/3 (all evidently a part of the same collection, India, Klein)^ 
and these other sheets are large plants which show that P. aUerni- 
folium Willd, perhaps equals Phymatodes hanerjiana Pal & Pal 
(Amer. Fern Journ. 53: 103. 1963), a species only recently segre- 
gated from P. scolopendria. I pointed this out to Dr. Pal, but he 
prefers to keep his species hanerjiana separate (I.e. lOS) on some 
characters that I can not follow. In any case, P. alternifoUum is 
different from P. nigrescejis; Presl's P. ylukenetii should therefore 
be placed as a synonym of P. nigrescens and not of P. alter nifoliunu 
Incidentally, there are plants in cultivation in the United States 
as P. scolopendria that appear to be surely P. alternifoUum, and 
such plants are to be expected in botanical gardens elsewhere. 
The characters of these species are well stated by Pal and Pal. 

Page 27. The new combination Pyrrosia costata (Presl) Holttum 
was previously made by Tagawa and Iwatsuki (Acta Phytotax. 
Geobot, 22: 100. May, 1967). Tagawa and Iwatsuki cite the 
wrong basionym Niphoholus costatus Presl ex Beddome (Ferns 
Brit, Ind. ad L 120. 1866, excL tab.), which is a later use of the 
epithet costatus but which is based ultimately on the same type 
as Apalophlebia costata Presl. All these names are based on Poly- 
podium costatum Wall. List no. 265. 1828, noni. nud. 

Page 30, Blechnopsis malaccensis Presl is referred to Blechnum 
indicum Burm., which should rather be P. serrulatum L. C. Richard. 
In the 'Tndex Filicum, Suppl. 3" Christensen definitely equated P. 
indicum Burm., previously regarded as dubious, with the common 
and widespread B. serrulatum, but the basis for this opinion is 
unknown. An examination of the type of B. indicum Burm. in 


serrulatum, but is, strangely enough, Asplenium longissimum 


cation I can confirm. Fortunately the well-known name Asplenium 
longissimum Blume need not be renamed ''A. indicum'' because 
there exists (but only very recently!) an Asplenium indicum 

122 American Fern Journal 

Sledge (Bull Brit. Mus. Nat, Hist. 3: 264, 1965), an entirely 
different plant. 

Page 33. Diplazium vestitum Presl. There were two syntypes 
cited— Cuming 333 and 336. Holttum cites 336 as the ^'Type/' 
but this should be ''Lectotype." He stated that he did not find 333 
at Prague, which is probably to be explained by PresPs comment 
(Epim. Bot. 84. 1851) under his D. grammitoideSy ''Affints sed diversa 
species est D. tenerum ex insula Leyte {Cuming n, 333) J^ This indi- 
cates either that Cuming 333 was a mixture in PresPs Herbarium, 
one sheet being Z). vestitum and one an undescribed species that he 
called ''D. tenerum/^ or that Presl changed his mind about the 
status of 333. Since Holttum did not find 333 filed as D. vestitum^ 
it is likely that the latter is true, and that 333 is filed as ''D. tene- 
rum. ^^ Holttum states that 333 is not represented at Kew; however, 
it is there, filed as D. ''sylvaticum,'' (Morton photograph 18620) 
and I have found it also in Firenze (Morton photograph 16124). 
A peculiar thing is that J. Smith cited 333 as from Samar, and the 
Kew and Firenze sheets are from Samar and also the specimen 
cited by Presl as his D, vestitum, whereas the specimen mentioned 
by Presl as '*D. tenerum'' was said to be from Leyte, which was 
probably an error, since the same number was not assigned to 
Cuming collections coming from different islands even though the 
plants might have been considered the same species. The speci- 
mens of Cuming 333 that I have seen are apparently referable to 
D, petiolare Presl and not to D. vestitum, which is more divided 
and more scaly on the rhachis. Cuming 336 was fixed as lectotype of 
D. vestitum by Hooker (Second Cent. Ferns t. 46, 1860 and Sp. 
Fil. 3:260. 1860), who referred only Cuming 336 to Asplenium 

vestitum (Presl) Hooker, and cited 333 under A. sijlvaticum (/. c 

248. 1860). 

Page 51. In his early and valuable work on Stenochlaena (Card, 
Bull. Str. Settl. 5:259. 1932), Holttum identified a rather common 



the type or the original description, which is quite understandable, 


Recent Fern Litekature 123 

from Singapore, where Holttum was working at the time. Now 
on seeing Presl's type (Cuming 226) he is certain that S. laurifolia 
Presl is a synonym of the common S. palustris (Burm.) Bedd., 
and that the species that he called ''S. laurifolia'' in 1932 needs a 
name. He remedies this by proposing ^'Stenochlaena cumingii 
Holttum, nom. nov. — S, laurifolia (non Presl) Holttum, Gard. 
Bull. Str. Settl. 5: (1932) 259; Copel, Fern Fl, Philip. 428." But 
this is an entirely inadmissible procedure, for one can give a new 
name only to a validly published species that is in need of a new 
name; a misidentification is not a vaUdly published species. This is 
not a new name but a new species, and as such it requires a Latin 
diagnosis and a designation of a type, neither of which Holttum 
provides. Therefore, S. cumingii Holtt. is a nonien nudum, not 
validly published. Dr. Holttum should provide a proper descrip- 
tion and a designation of a type to validate this name, which is 
needed.— C.V.IM. 


A. C. 

L. E. Schelpe. Contributions from the Bolus Herbarium, No. 1, 
published by The Bolus Herbarium, University of Cape Town, 
Kondebosch C. P., Republic of South Africa, pp. 1-132. 1969. 
Price 2 Rands. — There are at present no general treatments of the 
ferns of tropical Africa, although there are some fine regional 
treatments, such as :Madame Tardieu's "Les Pteridophytes de 
I'Afrique Intertropical Francaise," (1953), pteridophytes in the 
"Flore du Gabon," (No. 8, 1964), and "Flore de Cameroun," 
(No. 3, 1964), and Alston's "The Ferns and Fern-Allies of West 
Tropical Africa," (Suppl. ed. 2, The Flora of West Tropical Africa, 
1959). East tropical Africa has been unworked, except for the 
papers on Ethiopia by Pichi-Sermolli, and so Dr. Schelpe's new 
treatment will be of great value to students everywhere. This 
first part discusses the families Granmiitidaceae, Azollaceae, 
Salviniaceae, Vittariaceae, Lomariopsidaeeae, Adiantaceae, and 
PoljT)odiaceae (sens, restr.). The Grammitidaceae are considered 
to contain two genera, Grammiti^ and Xiphopteris (incl. Cteno- 
pteris); my union of these into the single genus is termed by 
Schelpe "a rather extreme standpoint," although Schelpe separates 

124 American Fern Journal 

these genera only by the character * 'Frond simple, entire to 
shallowly crenate" as opposed to 'Trond pinnatifid to deeply 
pinnatifid/' and adduces no other characters; there are in fact no 
other characters. Since I do not know of any other fern genera 
that are separated only by the character of blade division, and that 
only a difference between blades shallowly crenate as opposed to 
pinnatifid, my viewpoint does not seem unduly extreme. In fact, 
many fern genera that have blades normally pinnatifid to com- 
pound have some species wdth simple blades, e.g. Adiantum, 
TrichomaneSj Lindsaea, Polystichum, Thelypteris, Asplenium, 
Poly-podium, in fact many large genera. 

The new combination Xiphopteris alhobrunnea (Baker) Schelpe 
is proposed, based on Polypodium albohrunneum Baker (1877), but 
Polypodium sechellarum Baker (1874) is prior. Schelpe rejects the 
latter on the ground that it w^as based on two elements, one of the 
syntypes representing Z. albobrunnea and the other X. villo- 
sissima subsp. suhpinnata. Schelpe also states that the original 
description is not conclusive as to which element should bear 
the name sechellarum, and so rejects the name for either. This is 
contrary to the Code and to usual taxonomic procedures, which 
provide the workers should select lectotypes where these have not 
been provided. These lectotypes can be and frequently are quite 
arbitrary, where there is no special reason to select one rather than 
another element. But here there is a clear choice, since one of the 
syntypes is from the Seychelles Islands and the other from ]\Iauri- 
tius; since Baker chose the specific epithet ''sechellarum'' he ob- 
viously had this Seychelles specimen in mind, and so it should be 



replaces alhobrunnea. The following new combination is thus 


sechellarum Baker in Hook. & Bak. Syn. Fil. ed. 2, 508. 1874). 
Those who wish to continue to recognize Xiphopteris as a distinct 



I.ukungo River, Congo, but as I pointed out (Contr. U. S. Nat. 

Recent Fern Literature 125 

Herb. 38: 75. 1967) this species has been misidentified by recent 
authors; the type in the Willdenow Herbarium is clearly a synonym 
of >S. auriculata Aubl. I am not quite sure of the proper name for 
the species that has been called S, rotundijolia by Weatherby and 
others, but it may be S. minima Baker. Schelpe also records S, 
auriculata Aubl. from Lake Kariba, where it is a serious pest; 
however^ I have had an intimation from workers who are studying 
this problem that the Lake Kariba plant is not really yl. auriculata, 
but a sterile pentaploid, which would indicate that it is probably 
a recent hybrid that is spreading solely by vegetative means. 

Schelpe's treatment of Elaphoglossum is going to be especially 
useful, because this is one of the most difficult genera throughout 
the tropics. The key is based almost wholly on scale characters, 
which is doubtless inevitable, since the species are so uniform in 
most ways other than the scales. Recent studies of the stipe anat- 
omy and spores indicate that there may be some additional usable 

It is an open question w^hether Pteris quadriaurita Retz. is a 
species or an aggregate of many nearly allied species. Schelpe 
indicates that the question is not solved; in his treatment he is 
conservative and recognizes only one species, with four subspecies, 
which are not keyed out. Cheilanthes, one of the larger genera 
treated, is regarded in an inclusive sense, including Notholaena 
and AleuritopteriSj although Aspidotis is segregated, but Poly- 
podium is split up, following Ching's lead, into Pleopellis, Phyma- 
todes, Microgramma, Microsorium, and Polypodium. 

It should be mentioned that Schelpe recognizes Phymatodes K. 
B. Presl (Tent. Pterid, 195. 1S36) as a valid genus, but in pub- 
lishing this generic name Presl cited as synonyms Dipteris Reinw. 
and Microsorium Link, both earlier vahdly published and legiti- 
mate genera. Therefore, by Art. 69 of the Code, Phymatodes 
Presl w^as a superfluous name, and consequently illegitimate, smce 
the earlier name Dipteris should have been adopted instead of a new 
generic name. Phymatodes is, by Art. 7, Note 4, typified by the 
type of the name that ought to have been adopted, namely by 
Dipteris conjugata; it is a straight synonym of Dipteris, The name 

126 American Fern Journal 

Phymatodes could be used in Schelpe's sense by conservation^ but 
this hardly seems necessary- Its group is a small one, and is 
usually united with Microsoriumj even by those who split up 
Polypodium, such as Copeland, who is by no means conservative 
in his recognition of genera in this group. Schelpe separates 
Phymatodes from Microsorium by the former having paraphyses 
in the sori and the latter not, but even if true this is not necessarily 
a generic character, and that it is always true remains to be dem- 
onstrated. Schelpe had only a single species of Phymatodes and 
only two, closely allied, of Microsorium, The numerous Malaysian 
species of these groups have hardly been investigated. 

Schelpe uses the spelling ^^Microsoriumy^^ as is usual and correct 
in my opinion. The name was originally spelled '^Microsorum^^ 
by Link (Hort. Berol. 2: 110. 1833), and this spelling was adopted 
by Sledge in his treatment of the Polypodiaceae of Ceylon, but 
it was later changed by Link himself (Fil. Sp. 116, 135, 1841) to 
Microsorium. The Greek *'soros," generally Latinized as **sorus," 
is masculine, and if Link had wanted to make a generic name from 
it he would have called it ''Microsorus,'' just as he proposed 
Cyclosorus and Campfosorus. Instead, he evidently intended a 
diminutive, the Greek ''-ion," Latinized usually as ''ium,'* on the 
analogy of Pleridium, and so the original ending ''um" was properly 
corrected to ''ium/' — C.V.M. 

A New Na:me for a Species of Polypodium from North- 
western North America, by Frank A. Lang. Madrono 20: 53- 


60, 1969. — Lang has investigated the western Poly podium hesperium 
Maxon cytologically and finds that there are two cytotypes, a 
diploid and a tetraploid, which can be distinguished morphologic- 
ally. The type of P, hesperium corresponds to the tetraploid; it 
has the sori oval and medial, the rhizome with a sweetish, licorice 
taste, and the rhizome scales without a dark median stripe. It oc- 
curs mostly in the interior, east of the Cascade Mountains. The 
diploid has the sori circular and submarginal, the rhizome acrid, 
and the rhizome scales often with a median dark stripe. It occurs 
mostly in the western mountains from British Columbia to the 
Sierra Nevada, but there are a few specimens from elsewhere 

Recent Fern Literature 127 

(Wyoming, Colorado, Arizona). This diploid plant is called ''Poly- 
podium montense F. A. Lang, nom. nov.," and Pohjpodmm amor- 
phum Suksd. Werdenda 1 : 16. 1927, is cited as a synonym (Type: 
>Sfwfcsdor/ii^^7). The holotype of P. mo7?^ense is said to be Lay^g 211. 
This shows a confused concept, for if P. montense is a "nom. nov." 
I.e. a new name for a species previously published then it must have 



Latin description and cites a new and different type, his species is 
apparently really considered a new species. However, it is in my 
opinion a superfluous name, since the legitimate species P. amor- 
phum Suksd. is cited as a synonym and that name should have 
been adopted. Lang's reason for rejecting P. arnorphum is that he 
considers it a monstrosity to be rejected under Art. 71 of the 
Code. He defines a monstrosity as ''a plant that deviates greatly 
from the natural form of character, is abnormal, or is malformed," 
but this is by no means what the Code means by a monstrosity. 
A true monstrosity would be something like Spondylantha Presl, 
which was based on a plant of Cissus sicyoides that was so trans- 
formed by a smut that it was not recognizable as a Cissus at all. 
But Poljjpodium amorphum is not like that at all: it merelv has 
somewhat abnormal leaves but they are recognizable as belonging 


Many species have been described from somewhat abnormal speci- 
mens, e.g., Polypodium trijurcatum L., which was based on a plant 
with the fronds forked at the apex; the name obviously applies 
to a common West Indian species. So far as I know no plant with 
the leaves forked at the tip has ever again been found, yet no one 
has ever questioned the validity and legitimacy of the name P. 
irifurcatum. Therefore, the name P. moiiteme must be rejected 
as a superfluous new^ name for P. amorphum Suksdorf.— C.V.M. 

Flora Pal.\estina, Part one, Text, by Michael Zohary. 
Israel Academy of Sciences and Humanities, Jerusalem, 364 pp. 
+ 2 maps. 1966.— This flora, which includes the pteridophytes 
of Israel, is not new but has only recently come to my attention. 
As might be imagined from its aridity, Israel is not a "fernj" 


128 American Fern Journal 

country. In fact, there are only 15 species of pteridophytes that 
grow there. Any Fern Society members visiting Palestine will be 
able to identify the ferns they see merely by recognizing the 
genera, for each genus is represented usually by only one species. 
As a matter of fact, the families are so finely split that there is 
mostly only one species in each. The species are: Equisetum tel- 
mateiaj E, ramosissimumj Ophioglossum lusitanicum ^ Cheilanthes 
fragrans, C. catanensis {Notholaena vellea), Adiantum capillus- 
veneris, Pteris vittataj Anogramma leptophylla, Thelypteris pains- 
trisj Asplenium acUafitum-nigrum subsp. onoptens, Ceterach offici- 
naruniy Phyllitis sagittata, Dryopteris viUarii subsp. anstralisy 
Polypodium vulgare var. serratum, and Marsilea minula. — C.V.M. 

American Fern Society 

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Vol. 60 October-December, 1970 No. 4 

American Jfern S^ournal 


1Piibli«h«l by «fa* 









A Major North American Range Extension for the Forked Spleenwort, 
Asplenium septentrionale ■* • • David L. Emory 129 * 

Hard Water as a Limiting Factor in the Distribution of IsoEtes echino- 
Bpora LeRoy Lke and Filvnces Belknap 134 



The Rhizome Scales of Platycerium Barbara Joe Hoshizaki 144 

Shorter Notes: Stem Oross-section Prints Aid in Identifying Horsetails; 

•Psilotum nudum Spreading Northward ; -Terminology of the Spor- 

angial Structures of Equisetum ; Kinds of Cystopteris in California 161 

Notes and News 

Recent Fern Literature 

American Fern Society 

Index to Volume 60 

Errata ^■•. 







^ ' +'f ^•^ j{\M :01Q71 

®l)e American jFern feotietp 

Connctl foe 1970 

Wakren H. Waqnbb. Jr., Department of Botany, University of Michigan, 
Ann Arbor, Michigan 48104. Presidmi 

JoHM T. MiCKEii, New York Botanical Garden, Bronx Park, Bronx, New 
York. 10458. Vice-President 

A. MuBRAT Evans, Department of Botany, University of Tennessee. Knox- 
vifle, Tenn. 37916. Secretary 

LeRot K. Henry, Division of Plants, Carnegie Museum, Pittsburgh, Penn- 
sylvania 15213. Treasurer 

David B. Lellinoer, Smithsonian Institution, Washington, D. C 20560. 


Rational ^octetp B&epretfentatibeK 

Warren H. Wagner, Jr., University of Michigan A.AA.S. Representative 
Roll A M. Trton, Jr., Harvard University AJ.B.S. Representative 

American jf etn STournal 


10 B. Lellinger Smithsonian Institution, Washingt- 

\ Morton Smithsonian Institution, Washingt 

LA M. Tryon, Jr. 

Gray Herbarium, Harvard University, Cambrid; 
L. Wiggins.^ ^.Dudley Herbarium, Stanford Universitv, 


An illustrated quarterly devoted to the general study of ferns, owned by 
the American Fern Society, and published at 3110 Elm Ave., Baltimore j 
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Matter for publication should be addr^sed to the Editor-in-Chief. 

Changes of address, applications for membership, subscriptions, orders 
for back numbers should be addressed to the Treasurer. 

Subscriptions $5.50 gross, $5.00 net (agency fee $0.50) ; sent free to mem- 
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American Jfern Journal 

Vol. 60 October-December, 1970 No. i 

A Major North American Range Extension for the 
Forked Spleenwort, Asplenium septentrionale 

David L. Emory^ 

The ''Forked Spleenwort," Asplenhim septentrionale (L.) Hoffm., 
is a very unusual Spleenwort and one of the most striking ferns in 
the American flora. Presumably because of its peculiarly attenuated 
leaves (Fig. 1), the plant was recently referred to by Darling, 
(1964, p. 200) as an ''oddity/' Nevertheless it is a very wide ranging 
plant. Its distribution in western North America has been sum- 
marized by Lang (1969), who found the species recently for the 
first time in Oregon. Because of its ''striking resemblance to tufts 
of grass" and its tendency to occur in very localized populations, 
it probably occurs in other as yet undiscovered places in the 
western United States; at present it is known there from the eastern 
slope of the Rockies from South Dakota and AVyoming south to 
western Oklahoma, New ^Mexico, Arizona, and Baja CaUfornia. 
In the Old World, Broun (1938) reports it from the British Isles, 
Europe, northern Asia, and the Himalayas. Of the seven species 
of Spleenworts known in Britain, IManton (1950, p. 98) writes 
that "The rarest is Asplenium septentrionale . . ., a plant of south- 
ern affinities only found in a few presumably relict localities in the 
mountains of England, Scotland, and Wales," The habitat given 
by Broun is "sheltered crevices in cliffs of igneous rocks." 

During the second term of the 1970 summer session at :Mountain 
Lake Biological Station of the University of Virginia, I was enrolled 
in the Pteridology course which has been taught there smce 1961 
by Dr. Warren H. Wagner, Jr., of the University of :\Iichigan. A 
major emphasis in this course is upon student research in the field 
and laboratory. Accordingly, on August 2, 1970, a committee from 

^ I wish to thank Mrs. Carolyn Crump for the photographs and Dr. Florence 
S. Wagner (working under NSF Project GB-8113) for the chromosome obser- 

Volume 60, No. 3, of the Journal, pp. S9-12S, was issued Sept. 16, 1970. 

American Fern Journal 

Volume 60, Plate 16 



- ^ -^^^^r-.^ Wl^*--\^?yW-"' " "■" '^ ^" "-" V*'.\^-V-"' '^-'-'J^i j.T_rV 1 r r^ *■ - .x^^Vh-^^^v j'.T^> 

■-^iJ t_ >>*i^ j_^>** t --■ 

Bluff in Monroe Co., West VmamiA, where A. septentrionale Grows 

Range Extension for Aspleniu.m 131 

the class, consisting of Carolyn Crump, Ronald and Shirley 
Fortney, Lewis JNIorgan, and me, began a pteridological survey of 
Alonroe County, in southeastern West Virginia. This county, 
which is adjacent to Giles County, Virginia (the location of the 
Biological Station), seemed especially promising for field explora- 
tion because the class had already visited locations there discovered 
by E. T. Wherry durhig the late 1930's for Phegopteris connedilis 
and Lycopodium porophilum, which are growing at this southern 
latitude at unusually low elevations for these northern plants. 

While we were climbing on the upper slopes of a bluff of loose, 
strongly brittle shale above a stream {Plate 16), we noticed what 
appeared to be several tufts of a small grass or sedge growing from 
crevices in the exposed, brightly sunlit, and extremely dry shale. 
In color, size, and texture, the narrow leaf tufts also seemed quite 
similar to bunches of pine needles. Closer examination revealed, 
however, that these ^'needles," some of which were four inches 
long, were actually slightly expanded near their tips and forked. 
They bore on their undersides unmistakable fern sporangia. Hav- 
ing seen Asplenium septentriojiale on the American Fern Society 
Colorado Foray in 1964, I quickly realized that we had made a 

remarkable discoverv. Consultation of the literature indicated that 
our plants were nearly 1,200 miles from the nearest known station 
in western Oklahoma. 

When Dr. Wagner and the entire class visited the area on the 
following day, we counted sixty plants from four to nearly 40 
feet above the stream level on the northwest-facing bluff. Diligent 
searching up and down the stream revealed five more plants on 
another cliff about one-fifth of a mile upstream. One of the latter 
plants is shown in Figure 1, growing from the same crevice as A, 
plat y neuron. 

As these plants on both cliffs were of all ages, the populations 
are definitely reproducing. Unless some calamitous disturbance 
takes place, the ferns should be able to maintain themselves 
indefinitely. We have no idea how old the populations are. It is 
entirely possible that other stations will be discovered, as there 
are numerous similar habitats nearby. 

American Fern Journal 

Volume 60, Plate 17 

^,,^_._.^^^j,j-^j._-_--j,.r.-j,^^^jgy^^-, --_-^^-^-,-^.,-^^. r-^ 


■ HihF_V^|H~'Cri 

Fig. 1. AsPLExiUM j 


AS A. platyneuron (below). Fig. 2. Chromosomes of A. septentrionale 


Raxge Extension for Aspleniuai 133 

Because of its great disjunction and the possibility that the local 
populations have become differentiated, we were especially inter- 
ested in determining the chromosome number. Alanton (1950. p. 
98ff.) reported that A. septentrionale in Great Britain is tetraploid 
{n = 72 pairs at diakinesis). Cytological preparations of the plants 
we obtained; made by Dr. Florence S. Wagner, showed unmistake- 
ably the same number {Figs. 2^ 3), 

The * 'Septentrionale Bluff/' at an elevation of about 2100 feet, 
is shown in Plate 16. It supports only a sparse vegetation. Trees 
include Pinus virginiana^ P. strohus, Quercus prbiuSj Q. ilicifolia, 
Betula lenta, and Robinia pseudoacacia. Among the shrubs are 
Vaccinium vacillans, Kalmia latifoUa, Rhus rach'cans, and Parthe- 
nocissus quinquefolia. The herbs Aquilegia canadensis^ Houstonia 
longifolia, and the attractive Campanida divaricata are in evidence 
here, the last in bloom at the time of our discovery of the fern. 
The loose shale is extensively covered by a variety of lichens 
(especially such fruticose types as Cladonia ■mngiferina) and a few 
mosses, including Polytrichum spp. and Leucohryum glaucum. 

Ferns associated with the Forked Spleenwort are Asplenium 
trichomanes and A. platyneuron. In Great Britian and Europe A, 
septentrionale hybridizes with the first to yield A. X germamcum 
(Manton, 1950, p. lOOff.), This is the first time that A. septentrio- 
nale and A, platy neuron have been reported growing together. No 
hybrid plants of the latter parents have yet been discovered. 

A number of fronds collected from several plants on both days 
of our visits (Emory 70019, 70020) have been distributed to the 
following herbaria: US, GH, NY, MICH, WVA, and TENX, 
More precise information on the location in Monroe County, West 
Virgmia, is not being pubUshed in the hope that the population 
can be protected. I do not feel that there is a need for oversampling, 
and I wish to urge the greatest caution in maintaining these plants 
in their natural state without disturbance. 

This report ol Asplenium septentrionale in the Appalachian region 
constitutes a new addition to the flora of the eastern United States. 
The significance of disjunct eastern locaUties for western fern 
species will be discussed in another paper. 

134 American Feen Journal 

Literature Cited 

Broun, M. 1938. Index to North American ferns. 217 pp. Science Press, 

Lancaster, Pa. 
Darling, T. 1964. Southwest Vacation. Amer. Fern J. 54: 197-205. 
Lang, F. A., 1969. The first record of Asplenium septentrionale (L.) Hoffm. 

in Oregon. Amer. Fern J. 59 : 45-47. 
Manton, Irene. 1950. Problems of Cytology and EvohUion in the Pterido- 

phyta. xi +316 pp. Cambridge Univ. Press. 

jNIercersburg Academy, MercersbijrGj Pennsylvania 17236 

Hard Water as a Limiting Factor in the 
Distribution of Isoetes echinospora 

LbRoy Lee and Frances Belknap 

Isoetes in Wisconsin has been shown by Swindale and Curtis 
(1957) to be a typical member of the soft water flora. Tryon et al. 
(1953) show it distributed in soft water areas, but absent from 
areas where hard water lakes are common. Moyle (1945) reports 
the Quillworts in JMinnesota as members of a flora limited to a 
single chemical type: the soft water lakes found in the northeastern 
portion of the state. 

Although soft water is the typical location of Quillworts, they 
have been found in hard water. Lee reports Isoetes echinospora 
from Moshawquit lake. This Wisconsin lake has a specific con- 
ductance of 173 micromhos/cm and a total alkalinity of 98 mgm/1 
CaCOa, which exceeds the upper limit of tolerance as reported 
by Moyle (1945). Seddon (1965) reports it in two hard water lakes 
in Wales, Llyn Llygerian and Llyn Llywenan. 

:\Ioyle (1945) suggests that the aquatic species of Isoetes are 
found in soft water because they can not tolerate a total alkalinity 

Distribution of Lsoetes 135 

of above 50 mgm/1 CaCOs, but since they have been report eJ 
from other than the usual soft water sites, the distribution can not 
be completely attributed to water chemistry. Seddon (1965) con- 
cluded from field observations that competition is responsible for 
the restriction of I. echinospora to soft water. 

To determine if Quillworts could survive and grow in hard 
water without competition, /. echinospora var. hraunii (Dur.) 
Engelm. was grown under laboratory conditions of varying water 
chemistry during one summer growth period. 

Table I, Water Analysis of Experimental Tanks^ 

alkaUniiy . conductance water color 

(CaCOi mgm/l) pH {micromhos/cvi) (Pl-Co units) 

Crooked Lake 29 7.5 119 15 

Crystal Lake 92 7.6 231 35 

Wis. River 112 8.2 282 65 

Fish Lake 148 7.9 304 35 

Pretty Lake 160 8.3 483 40 

Lake Ripley 184 8.6 431 20 

LakeWingra 228 7.8 643 38 

Merrill Springs 281 8.0 585 30 

^ Water analysis furnished by Mr. Ronald Poff, Water Classification Special- 
ist, Wisconsin Department of Natural Resources, Madison, Wisconsin. 

Eight 10-gallon aquaria were filled with water and substrate, each 


a range from very soft to very hard water (see Table I), Experimen- 


For each tank, ten plants were randomly selected from the collec- 


then planted. Twelve hours of light was supplied daily by Gro-lux 
fluorescent bulbs to provide 100 ft-c of illumination at the water 
surface. Weekly water analyses determined that little change in 
water chemistry occured over the experimental period. 

136 A:\iERicAN Fern Journal 


At the end of 21 weeks all plants were still surviving in the 
soft water control (Crooked Lake) and the hard water aquarium 
from Ripley. Seventy per cent of all the experimental plants 
survived; they grew an average of 5 new leaves and had 2 cm of 
leaf and root growth, the same as the average in the soft water 
control tank, Sporulation did not occur. 

At the termmation of the experiment the plants were replanted, 
additional hard tap water (280 mgm/1 CaCOa) was added to each 
tank to replace evaporation loss, and continuous illumination at 
a high intensity was provided. After five months all plants still 
survived and appeared healthy. Several were sporulating. 

While the results of this experiment are tenuous due to the death 
of 30% of the experimental plants, they do support the behef that 
water hardness is not the single major limiting factor in distribu- 
tion. The continued growth and sporulation with increased illumi- 
nation, plus the fact that the tw^o tanks which had 100% survival 
were significantly less colored than the others (see Table I), 
indicates that the amount of light reaching the plants may be an 
important factor in the distribution of Isoetes ecliinospora var. 

Literature Cited 

MoYLE, J. 1945. Some chemical factors influencing the distribution of aquatic 

plants in Minnesota. Amer. Midi. Nat. 34: 402-420. 
Seddon, B. 1965. Occurrence of Isoetes echinospora in eutrophic lakes in 

Wales. Ecolo©^ 46: 747-74S. 
SwiNDALE, D. and J. Curtis. 1957. Phytosociology of the larger submerged 

plants in Wisconsin lakes. Ecology 38 : 397-407. 
Tryon, R., N. Fassett,-D. Dunlap, and M. Diemer. 1953. The ferns and 

fern allies of Wisconsin. 158 pp. University of Wis. Press, Madison, 


Madison Public Schools, IMadison, Wisconsin 53715. 

Pteridophyte Folk Remedies 137 

Indian Pteridoph3rtes Used in Folk Remedies 

H. S. PuRi^ 


The medicinal qualities of ferns^ real or imaginary^ are mentioned 
as early as 300 B.C. by the Greek philosopher Theophrastus (Corne, 
1924a) and by his Indian contemporaries Sushrut and Charak. 
Dioscorides also speaks of brake fern^ male fern, and others. Among 


the medicinal plants mentioned in old Indian texts, two are def- 



meaning peacock^s feather^ has been identified as Adia7}tii7n cau- 
datum L. by some persons; others consider it to be Adiniopteris 
radiata (Swartz) Link. The other drug called ^'Hanspad*' or 
^'Hansraj/' meaning swan's foot, appears to be some species of 
Adiantujn, most probably A. capillus-veneris L., A. lumdatum 
Burm. f. or i4. vemistum D. Don. In addition to these plants there 
are about 40 species of pteridophytes used as drugs. Of these 
Dryopferis filix-mas (L.)'Schott is included in the pharmacopoeias 
of many countries. In the 1955 Pharmacopoeia of India this plant 
is replaced by the allied species, Z). odontoloma (Moore) C, Chr., 

r ^^ 

D. marginata (Wall.) Christ, and D. schimperiana (Hochst.) C. 
Chr., and in the 1966 Pharmacopoeia of India by D. odontoloma, D. 
marginata^ D. chrysocoma (Christ) C, Chr,, D. ramosa (Hope) C. 
Chr., and D. harhigera (Hook.) Ivuntze. Lycopodium davatum is 
included in the Indian Pharmaceutical Codex (Mukerji, 1953) and 
in the proposed Indian Homeopathic Pharmacopoeia. Equisetum 
arvense L. is included in the U.S.S.R. Pharmacopoeia (1961) and 
in the German Pharmacopoeia (Wealth of India); and Puri (1969) 
has recommended its inclusion in the Indian Pharmacopoeia. The 
species oi Adiantum mentioned above are used unofficially in some 
of the patented cough preparations marketed in India- The rest 
of the plants mentioned below are used in domestic and rural 

1 The author is greatly indebted to Prof. P. N. Mehra for his keen interest 
d to Mr. C. V. Morton for his kind suggestions during the preparation 


of this article. 

138 American Fern Journal 

A diniopteris radiata (Swartz) Link.— Used as an anthelmintic 
(Watt, 1889; Kirtikar & Basu, 1935; Chopra et al, 1958; Chunekar 
& Pandey, 1969), as an alterative in prolonged malarial fevers 
(Nadkarni, 1954; Chopra et al.), and as an astringent to arrest 
hemorrhages (Chunekar & Pandey). 

Adiantum aethiopicum L. — Infusion of leaves used as an emol- 
lient in coughs and diseases of the chest (Caius, 1935). 




the fronds of this plant is considered quite effective in all types 
of bronchial troubles (Watt; Nadkarni; Steinmetz, 1954; Wren, 
1956; Uphof, 1959). An hifusion of the herb serves as a shampoo 


dandruff and also promotes hair growth (Watt 


all types of fevers (Nadkarni; Biswas, 1955). It is used for hydro- 
phobia by the physicians of the Persian system of medicine (Watt). 

Adiantum caudatum L.— Fronds used for diabetes, coughs 
(Kirtikar & Basu; Chopra et al), and for migraine (Stewart, 
1869). Also used externally for skin diseases (Chopra et al). 

Adiantum lunulatum Burm. f.— One of the constituents of 
Hansraj, the drug esteemed in India for coughs. It is considered a 
bronchio-dilator, diuretic (Chopra et al) and pectoral (Caius). In 
western India used extensively in the treatment of fevers of 
children; the rootstock is considered good for fever and elephan- 
tiasis (Kirtikar & Basu). 

Adiantum venustum D. Don— According to Kirtikar and Basu 
the fronds of this plant are the source of the Indian drug Hansraj. 
Because of the black color of the stalks of the fronds, it is some- 
times known by the name of black Hansraj . The plant is adminis- 
tered as an anodyne in bronchitis and is considered a diuretic and 
emmenagogue (Stewart, Ku-tikar & Basu; Chopra et al). It is a 
tonic, febrifuge, and expectorant, and is particularly useful as a 
tonic during convalescence from fevers (Caius). 

Asplenium adiantum-nigrum L.— A decoction or syrup of the 
fronds is used as an emmenagogue (Kirtikar & Basu). It is bitter, 

Pteridophyte Folk Remedies 139 

diuretic, and laxative, and is considered useful in ophthalmia and 
diseases of the spleen and jaundice. It is believed to cause sterility 
in women (Chopra et al.). 

Asplenium falcatwn Lam. — Used in enlargement of the spleen, 
incontinence of urine, calculus, jaundice, and malaria (Kirtikar 
& Basu). 

Asplenium ruta-muraria L, — It is used for rickets (Chopra et 
al), and as an expectorant (Xadkarni) and deobstruent (Kirtikar 
& Basu). According to Steinmetz, ''Herba Rutae Murariae*' is 
an expectorant and cures diseases of the spleen. It prevents hair 
from falling and makes it grow (Fernie, 1914; Steinmetz), 

A splemum trichomanes L. — Considered a laxative (Fernie ; 
Chopra et al) and anthelmintic (Nadkarni). The leaves are 
sweet, mucilaginous, and are an expectorant highly useful in 
pulmonary disorders (Caius). 

Athyrhim filix'femina (L.) Roth. — Used as a substitute for the 
Male Fern, but has been found quite ineffective (Chopra et al). 

Botrychium spp. — Various species of this genus, such as B. 
liinaria (L.) Swartz, B. ternatum (Thunb.) Swartz (Chopra et al), 
and B. virgimamim (L.) Swartz (Kirtikar & Basu), are used in 
dysentery, ruptures, and for healing wounds (Wealth of India). 

Cibotium harometz (L.) J. Smith— The rhizome is used as a 
vermifuge and the root as a tonic. The long, silky, yellowish hairs 
at the base of the fronds cause rapid coagulation of the blood and 
when properly used are useful for arresting hemorrhages from 
capillaries (Wealth of India). The root is employed as a tonic in 
China, where it is said to exercise a special action on the genito- 
urinary organs (Kirtikar & Basu). Used in typhoid and in the 
treatment of dyspepsia and coughs (Chopra et al). 

Drynaria quercifoUa (L.) J, Smith.— The rhizome is bitter, 
tonic, and astringent to the bowels. It is also used in typhoid and 
in the treatment of dyspepsia and coughs (Chopra et al). A decoc- 
tion of the rhizome is used as an astringent (Uphof). 

Dryopteris fiUx-mas (L.) Schott.— One of the oldest anthelmin- 
tic druss, commonlv known as Male Fern, and used since ancient 

140 American Fern Journal 

times for expelling tape-worms from the intestines. This fern does 
not grow naturally in India, but various allied species which are 
up to British Pharmacopoeial and United States Pharmacopoeial 
standards are used (Indian Pharmacopoeia, 1955, 1966). The use 
of D. filix-mas does not appear to be harmless. According to 
Steinmetz, its application may cause paralysis of the muscles and 
nerves, and even blindness. This may be due to the presence of 
the enzyme thiaminase, which destroys thiamin (Pohl, 1955). 
Equisetum arvense L.— "Herba Equisiti Arvensis" has diuretic 



of blood in urine (Steinmetz). An oral administration produces a 
decided increase in blood corpuscles. The plant is used for acidity 
of the stomach and dyspepsia. (Wealth of India). 

^ Equisetum debile Roxb.— Diuretic and astringent (Biswas), and 
given for gonorrhoea (Stewart). 

Helminthostachys zeylanica (L.) Hook.— The plant has intoxi- 
cating and anodyne properties and is used in sciatica (Chopra et 
al; Wealth of India). Rhizome used for malaria (Uphof). The 
Malays regard the rhizome as a tonic, and eat it with betel for 
whooping cough. In Java used for dysentery, catarrh, and the 
early stages of phthisis (Wealth of India). 

Lycopodium cernuum L.—In Malaya a decoction of the plant 
used as a lotion in beri-beri and also for coughs and uneasiness in 
the chest. An embrocation of the ashes in vinegar is recommended 
for skin eruptions (Wealth of India). 

Lycopodium clavatum L.— Mainly the spores of this plant are 
used in medicine. English druggists included this powder in their 
list of drugs before 1692. It was introduced in the U.S. Pharma- 
copoeia of 1860 (Whitebread, 1941). This drug has been used 
mternally with good results in urinary disorders, such as the 

cat arrh al 


ureter, and 

and as a general and gastric sedative (Wren). The plant is also 


Pteridophyte Folk Remedies 141 

of a tincture it is given for rheumatism, epilepsy, and pulmonary 
disorders (Nadkarni). An aqueous extract of it has been employed 
in various countries as an antipyretic (Wealth of India, 1902). 

Sometimes the spores of L, annotiniim L., L. comjJanatum L. 
and L. selayo L. are also used for the above purposes (Wealth of 
India). Frequently LycojJodium spores are adulterated with such 
substances as slightly roasted and colored starch, dextrin, and 

various pollens (Whitebread). 

The Lycopodium powder has side effects. Whitebread has quoted 
instances where it has acted as foreign body and has given rise to 
an indolent chronic post-operative inflammatory reaction. The 
alkaloids of this drug have also been found harmful to warm- 
blooded animals. A variety of reactions, such as pressor effects, 
stimulation and contraction of the uterus, and paralysis are at- 
tributed to these alkaloids (Wealth of India). 

Lygodium circirmattim (Burm. f.) Swartz — The spikes of this 
plant are reported to be used in Indonesia as an external applica- 
tion for wounds (Wealth of India). 

Lygcdium flexuosum (L.) Swartz — Used as an expectorant. Fresh 
rootstocks are applied externally for rheumatism, sprains, scabies, 
eczema, and cut wounds, and are reported to be particularly useful 

for carbuncles (Caius). 

Lygodium japonicnm (Thunb.) Swartz— Used as an expectorant. 
In China a decoction of the vegetative parts and spores is used as 

a diuretic (Wealth of India). 

Lygodium microphyllum (Cav.) R. Brown— A decoction of leaves 
given for dysentery. Leaves are also applied as poultices for skin 
diseases and swellings (Kirtikar & Basu). 

Mamlea quadrifolia L— This is probably the plant used m 
ancient India under the name of *^Sparka" for leprosy and skin 
diseases, fever, and for poisoning of blood (Chunekar & Pandey). 

Ophioglossum vulgatum L.— The leaves boiled in oil or fat are 
considered a remedy for wounds. This preparation cools inflam- 
mation (Fernie; Chopra et al). 

Osmunda regalis L.— Considered a tonic and styptic, and used 
against rickets (Chopra et al). The middle part of the plant boiled 

142 American Fern Journal 

in liquid is considered good for wounded persons. The rootstock, 
stamped in water or gin until the liquor becomes a stiff mucilage, 
has been used to cure back pains (Fernie). 

Poly podium vulgare L. — An infusion of ''Herba Polypodii" is 
a mild laxative and expectorant. An aqueous extract of "Rhizoma 
Polypodii" is recommended in the obstruction of viscera; liver and 
gland diseases, and coughs and chills (Steinmetz). In northern 
India this or a related species is used as an alterative (Stewart). 

Polydichum squarrosum (D. Don) Fee— The rhizome of this 
plant is sold in the Indian market under the name ''Nirbissi/' 
and the drug is used as an antidote to poisons; it is a possible 
substitute for the male fern (Mittal, 1953). 

Pteridium aquilinum (L.) Kuhn. — The rhizome is astringent and 
is useful for diarrhea and inflammation of the gastric and mucous 
membranes. Boiled in oil or hog's fat, the rhizome is made into an 
ointment for wounds. A decoction of the rhizome and fronds has 
been given in chronic disorders arising from obstruction of the 
viscera and spleen (Wealth of India). It is also used instead of hops 
(Corne, 1924b). 

Pteris ensiformis Burm. f. — The juice of the young plant is 
stated to possess astringent properties; a decoction of fresh fronds 
is given in dysentery. The juice of the rhizome is applied in the 
glandular swelling of the neck (Wealth of India). 

Pteris multifida Poir. (P. serrulata L. f.)-In China a tincture or 
decoction of rhizome is given in dysentery. Also said to be a good 
vermifuge. The toasted fronds and rhizome are made into a paste 
with sesame oil and applied to the skin afflictions of infants 

(Wealth of India) . 

Selaginella spp. — Apparently dry plants of some species appear 
to become fresh and green again when put into water. Because of 
this, physicians of the Indian system of medicine use it as a re- 
juvenator for the human body. It is also given in mental diseases, 
rickets, blood vomiting, and spermatorrhoea (Chunekar & Pandey). 

Tectaria polymorpha (Wall.) Copel. — Used as an anthelmintic 
(Kirtikar & Basu). 

Pteridophyte Folk Remedies 143 

Literature cited 

Biswas, K. 1955. Common Medicinal Plants of Darjeeling and Sikkim. 

Botanic Garden, Calcutta. 
Caius, J. F. 1935. The Medicinal and Poisonous Ferns of India. J. Bombay 

Nat. Hist. Soc. 38: 341-361. 
Chopra, R. N., Chopra, I. C, Handa, K. L., and K\pur, L. D. 1958. Chopra's 

Indigenous Drugs of India. U. N. Dhur and Sons, Calcutta, 
Chunekar, K. C. and Pandey, G. S. 1969. Bhavparkash Nighantu (in Hindi). 

The Chowkhamba Vidya Bhawan, Varanasi, India. 
CoRNE, F. E. 1924a. Ferns— Facts and fancies about them — II. Amer. Fern 

J. 14: 77-82. 

1924b. Ibid.— III. Amer, Fern J. 14: 115-118. 

Fernie, W. T. 1914. Herbal Simples, ed. 3. John Wright & Sons, Bristol. 
KiRTiKAR, K. R, and Basu, B. D. 1935. Indian Medicinal Plants, Part IV, 

ed. 2. Lalit Mohan Basu, Allahabad, India. 
MiTTAL, T. C. 1953. Pharmacognosy and phytochemistry of some Indian 

species of Polystichum and Athyrium. M. Pharm. Thesis, Panjab 

Univ., Chandigarh, India. 
MuKERji, B. 1953. Indian Pharmaceutical Codex, vol I. Council of Scientific 

and Industrial Research, New Delhi. 
Nadkarni, K. M. 1954. Indian Materia Medica, vol I. Popular Book Depot, 

PoHL, R. W. 1955. Toxicity of Ferns and Equisetum. Amer. Fern J. 45 : 95-97. 
Pharmacopoeia of India. 1955. Manager of Publications, New Delhi. 

. 1966. Second edition. Manager of Publications, New Delhi. 

PuRi, H. S. 1969. Scope of including some indigenous medicinal plants in I. 

P. Indian J. Hosp. Pharm. 6: 102-106. 
State Pharmacopoeia of the Union of Soviet Socialist Republics. 196 L 

9th edition, Moscow. 
Steinmetz, E. F. 1954. Materia Medica Vegetabilis. E. F. Steinmetz, Kaizer- 

gracht, Amsterdam, Netherlands. 
Stewart, J. L. 1869. Panjab Plants. Gov't. Printing Press, Lahore. 
Uphof, J. C. Th. 1959. Dictionary of Economic Plants. Hafner, New York. 
Watt, G. 1889-1892. Dictionary of Economic Products of India, Parts I-IV. 

W. H. Allen. 

Wealth of India. 1948-1969. Parts I-VIII. Council of Scientific and Indus- 
trial Research, New Delhi, 

Whitebread, C. 1941. Beware of **Lycopodium.*' Amer, Fern J. 31; 100-102. 

Wren, R. C. 1956. Potter's New Cyclopaedia of Botanical Drug and Prepara- 
tion. Sir Issac Pitman and Sons, London. 

Botany Department, Panjab University, Cil\ndigarh, India. 

144 Americajn^ Fern Journal 

The Rhizome Scales of Platycerium^ 

Barbara Joe Hoshizaki 

Considerable diversity exists in the structure of fern rhizome 
scales. Although the more conspicuous features of scales have long 
been used in describing and identifying some groups of ferns, the 
more detailed aspects have generally been ignored. Modern mono- 
graphs and treatments on ferns have contributed to a better 
understanding of the value scales may have in the interpretation 
of evolutionary relationships. The object of this paper is to de- 
scribe some of the rhizome scale features of Platycerium that seem 


ning the evolutionary trends within the 
ures have been uointed out bv Wasner 

Before the rhizome scales could be studied for structures that 


first the probable evolution of the Platycerium species. For this, 
I used a variety of characters other than those of the rhizome 
scales. Reliable characters included stele types, sporangial arrange- 
ment, nature of the sporangial stalk, and completeness of the 
annulus. Only after I had arrived at the probable evolution of the 
species did I examine the rhizome scales in detail to determine 
how their features correlated with the patterns I had determined. 
The three main lines of evolution are summarized in Table I. 
The Afro-American line includes P. amlinum and two branches 
both starting from P. quadridichotomum. One branch ends in P. 
angolense and the other branch ends in P. elUsii. The sequence of 
species in the other two lines is essentially Imear, as listed. 

In selecting rhizome scales for study several factors must be 
kept in mind. The scales on individual plants vary, but species 
differences are usually consistent, providing the factors of the age 
of the plant, weathering, and selection of a typical scale from the 
sampHng are taken into consideration. Immature or small plants 


FelWhT*^** supported by a National Science Foundation Science Faculty 

Rhizo:me Scales of Platycerium 145 

generally have smaller, lighter, and fewer bicolorous scales bearing 
fewer haks than those on mature plants. On fully mature plants 
scales may have suffered damage by w^eathering and must be 
assessed accordingly. The best scales for study are those that are 
starting to darken on full grown plants. Even in perfect material 
there will be variations; the largest, darkest scales usually show 
the characters of the species more clearly than the narrower, 
smaller, lighter colored scales or fibrils. In regards to selection of 
trichomes for study, Carlquist (1961, p. 33) aptly states, "Each 
trichome type in each species shows some variability and thus each 
type should be studied as a population having extremes and a 
typical condition." 

The plants used in this study are from cultivation; for most 
the specific place of collection is not known. ^ Silhouettes of the 
scales were made by tracing photographs; drawings of the hairs 
were made with the aid of a microprojector. 

Morphology and Evolutionary Significance of Scales and Their Hairs 

Scale distrihution, — Rhizome scales are most conspicuous on the 
buds, but also cover the entire surface of the stout rhizome and 
extend up the phyllopodia occasionally to just beyond the abscis- 
sion layer. On the bud the many light colored scales are tightly 
overlapped and appressed to the surfaces. Progressing to the older 
parts of the rhizome, the scales become darker, less appressed, and 
may twist or curl to form a thick, chaffy covering; they are per- 


General s/mpe.— Each plant bears scales of various widths and 
lengths; the larger, darker scales are here considered the represen- 
tative scales of a species. For most species these are more or less 
narrowly triangular. However, the scales of P. andinum and P. 
angolense (Figs. lA, 15) are noticeably linear-triangular, and in P. 

* Messrs. Ed Franks, Rudy Ziesenhenne, Marcel Lecoufle, and John Roach 
very generously provided most of the plants and specimens for this stiidy, 
and I am greatly indebted to them for their many kindnesses. I wish to thank 
Miss Marie Sanchez for her expert advice in preparing the illustrations and 
also to acknowledge the assistance and many valuable suggestions given to 
me by Dr. Mildred E. Mathias. The major part of this work was done at tne 
University of California, Los Angeles, where facilities were kindly provided. 


American Fern Journal 

coronarium and P, ridleyi they are obovate {Figs, 29 ^ 30). All but 
P. coronarium and P. ridleyi have fibrils (very narrow scales) 
which also vary in width and length on the same plant; the nar- 
rowest are uniseriate flattened hairs and the widest are indistin- 

Table I. Species and Their Rhizome Scale Characters 


Character stales* 
= very specialized 
2 = specialized 



= generalized 




Afro-American Line 
P. andinum 

P. quadridichotomum 
P. stemaria 

P. angolense 

P. madagascariense 

P. vassei 

P. ellisii 

Javan-Australian Line 
P. willinckii 
P. veitchii 

P. bifurcatum 
P. hiUii 

Malayan-Asiatic Line 
P. wallichii 

P. holttumii 
P. wandae 
P. grande 
P. coronarium 
P. ridleyi 










present mediate papery 












Base non- 

(0) , suh- 





guishable from narrow scales. The almost universal presence of 
fibrils in the genus seems to indicate the ancestral type had fibrils 
which developed into progressively broader scales. It is generally 
accepted (Bower, 1923, pp. 201-205) that hairs preceded fibrils and 
that fibrils developed into scales. FoUowiner this line of reasoning, 

Rhizome Scales of Platvcerium 


narrow scales probably gave rise to broad scales. Extremely broad 
scales {Figs. 26-30) seem to be linked to the development of 
flabelloid margins. The scale length varies from 2 to 35 mm, the 
longest being those of P. icandae and the shortest those of P. 

Hairs or 














ha irs 



(0) , sub- 

none (0), 


none or 


none ((ij, 

^P-fl-^H ■ 













(1), mier- 

mme of 









(2), all 


(0), 1-2 





spec. (3) 


(1), 1 (2) 


fcroad (3) 



































tne rnizome. ine lexiure ui luc ^>^ax^^ x^ ^ — — ^ , 

grande scales are thin and papery and those of P. coronanum and 
P. ridleyi are also except at the center; this is probably a special- 
ized condition. 

148 American Fern Journal 

Scale apex. — Scale apices may be long attenuate to acute, to 
blunt, rounded or emarginate (Fiys. 2-5). Typically the attenuate 
apices end in a long, often gland-tipped filament. Acute apices 
have fewer and stouter cells in the filament; blunt apices are 
multicelluar and sometimes apiculate or emarginate. The apices 
may often be branched into a number of hairs or filaments. If 
it is true that scales have been derived from hairs, then the general- 
ized scale apex would be attenuate and filiform and the specialized 
apex blunt and multicellular. There would be a trend toward 
increasingly broad apices. There remains, however, the possibility 
that the extremely long attenuate apices of P. angolense and P. 
andinum may be a specialized condition. 

In the clear evolutionary line of the ^Malayan-Asiatic group 
there is a series from generalized to specialized apices. The inter- 
mediate state consists mostly of acute apices that are rarely fili- 
form. The Javan-Australian group has the generalized to inter- 
mediate state; the Afro-American group tends to the generalized. 

Scale base. — The scale base may be slightly narrowed, rounded, 
truncate, or subcordate to cordate (Figs. 9-13); however, most 
are unmodified. Most scales are sessile and attached by the length 
of their basal edge {Fig. 9), which may be a broadened area if 
the scale is bent and appressed to the rhizome {Fig. 10). Scales 
may also be subpeltate (pseudopeltate) to peltate. In all cases 
the scales interrupt the rhizome epidermis at their point of attach- 
ment. The subpeltate condition (Ff^/. 12), found on some scales of 
P. willinckii, P. veitchii, and P. bifurcatum, is often accompanied 
by a subcordate or cordate base; the attaching tissue is a long or 
short flange. Platycerium hillii and some forms of P. bifurcatum 
have weakly peltate scales with rounded bases, which are most 
frequently found on the phyllopodia and which tend to detach 
easily {Fig. 13). There is a definite stalk or short flange which 
attaches to the rhizome in a sUght epidermal depression. Peltate 
scales are generally considered more advanced than subpeltate or 
other unmodified ones (Bower 1923, p. 200). 

Scale margin.— The scale margin may be entire, or bear papillae 
or hairs, or be fiabelloid (see the separate discussions on hairs and 

Rhizome Scales of Platycerium 149 

flabelloid margin). Small, dark protuberances are sometimes seen 
on or near the walls of the marginal cells of P. grander F. icandae^ 
and P. hifurcatum. They seem to be persistent cell nuclei which 
have not deteriorated, for in the younger parts of the scale nuclear 
stains bring out obvious nuclei which if examined in progressively 
older cells move toward the wall and appear as the small dark 

Bicolorous scales. — The central middle to basal areas of the scale 
are marked by a stripe on at least some scales of most species. 
The stripe is composed of cells which have all thickly pigmented 
brown to black walls. A scattering of lighter colored cells with 
only their lateral walls thickened may be found among the darker 
cells. As there is considerable inconsistency in color and pattern 
of these stripes, it is difEcult to use these to evaluate phylogenetic 
trends. The stripe extends to or nearly to the apex in members of 


than in other species. 

Platycerium grande often lacks a well defined stripe, particularly 
on the larger scales of the phyllopodia. 

Scale development, — Rhizome scales begin their development 
from uniseriate hairs. These hairs divide longitudinally, trans- 
versely, and clinally at the thickened center to form the scale. 
For cell details see Figure 1, In non-peltate scales the enlargement 
of the base is by transverse and longitudinal cell division. Peltate 
scale development in P. hUlii often shows a cupping in or turning 
under of the basal auricles, which are eventually obliterated by 
thickening, enlargement, and overtopping growth of the whole 
basal area, resulting in a thick, rounded base {Fig. 13), The 
attaching cells that were between the auricles become submarginal 
and differentiate into a short flange or well-defined stalk. This 
scale development is similar to that illustrated for Pyrrosia (Nayar, 
1961, figs, 7-15, p. 166), except that there are no auricles. 

Hair distribution,— Hsiirs are found on all scales except those of 
P. irallichii. All others have hairs along the edge of the scales; 
the hairs are sparse near the basal edge. Platycerium UJurcatum 
and P. hillii tend to have long, tangled hairs near the apical 

mnrfTin nf fViP c/>o1p>: hnf fVtis is not alwavs consistent (Fig, A), 

American Fern Journal 

Volume 60, Plate 18 



ATYCERiim Rhizome Scales 



Fig. 4. P. hillii, acute apex. Fig. 5. P. grande, blunt apex. Fig. 6. P. 



Rhizome Scales of Platycerium 151 


e apex 

occasionally in P. angoIe?ise, and over much of the surface in P. 
eltisii and P. vassei (but more sparsely in the latter) ^ and on the 
face of the marginal cells in P. ridleyi. The dense marginal fringe 
of hairs in P, hoUtumUj P. ivandae, and P. granule scales {Figs. 
6-8) arises from marginal and submarginal cells of the flabelloid 
margin. Having the hairs consistently scattered over the surface 
of the scale or spreading to a submarginal position occurs in most 
advanced to moderately advanced species and is to be regarded 
as more specialized than having hairs limited to marginal cell 
edges. In the apparently related genus Pyrrosia, a similar distribu- 
tion of hairs is also found among the advanced or moderately 
advanced species (Nayar, 1965, p. 13). 

Hair types, — The hairs found on the rhizome scales fall into 
three intergrading types; glandular, ray, and basic. The glandular 
types are typically stout, round at the apex, and contain a dark, 
gland-like substance. The ray types appear much like the rays of 
the stellate hairs of the lamina; most are whitish and blunt to 
pointed at their apex, and generally flat, although some are conical 
and often a bit darker. Basic hairs are close to the ray type in 
appearance and color, but are t^T^^^l^y bent, twisted, or crooked, 
and not as flat and white as the ray type. Shorter basic hairs bear 
a close resemblance to the marginal papillae, whereas longer ones 
may be setiform. Typical glandular hairs are found in P. ridleyi 
(Fig, 30), typical ray types in P. vassei (Fig. iPs), and basic types 
in P. gra?id€ (Fig, 28b), Some basic types may blend into the ray 


are found 

line (Figs, 25-30). The Javan-Australian line (Figs, 21-24) and the 
more primitive members of the Afro-American Hne (Figs, 16-18) 
have mostly basic to glandular types. Basic to glandular inter- 
mediate hairs tend to have the contents of the cell collapsed, 
causing a distortion of the cells (Fig. 17). 

Origin of the hair types,— The presence of hau'-like papillae and 
their similarity to basic hairs suggest that basic hairs are derived 
from elongate papillae. This follows the dictum that trichomes 

American Fern journal. 







Platycerium Rhizome Scales and Hairs. Figs. 9-13. Rhizome scale 


VASSEI, broad ATT. 


BASE. FiGS. 14-30. Rhizome scales and hairs showing changes along 


Fig. 15. P. angolexse. Fig. 16. P. stemaria. Fig. 17. P. quadridichotomum. 

A^OLUME 60, Plate 19 

Fig. is. P. madagascariense. Fig. 19. P. vassei. a, Mozambique j b, ^Iada- 
gascar; c, cultivated. Fig. 20. P, ellisii. Figs. 21-24. Javan-Australtan 
LINE. Fig. 21. P. willinckii. Fig. 22. P. veitchii. Fig. 23. P. bifurcatum. 
Fig. 24. P. hillii. Figs. 25-30. Malayan-Asiatic line. Fig. 25. P. wal- 
ucHii. Fig. 26. P. holttumii. Fig. 27. P. waxdae. Fig. 28. P. grande. a, 


NARiuM. Fig. 30. P, rtdleyl The abbreviation is: m ^ marginal cellj^. 

154 American Fern Journal 

originate from papillate epidermal cells (Netolitzky cited in Carl- 
quist, 1961, p. 34). This hypothesis is supported by the presence 
in P. holttumii and P. grande of very short to long papillae inter- 
mingled with basic hairs, which are distinct only by the cell wall 
separating the hair cell from the trichoblast. 

Uniseriate hairs are generally regarded as the basic type from 
which many other variants are derived (Wagner, 1964, p. 90). 
The origin of the glandular hair is from the basic hair. There are 
all degrees of gradation between these two types. Also, w^ell defined 
glandular hairs appear only in the more advanced members of the 
]Malayan-Asiatic {Figs. 29, 30) and Afro-American {Figs, 14, 15, 
19j 20) lines and hence correlate with other advanced traits. The 
Javan-Austrahan line has retained the intermediate basic to 
glandular hairs {Figs. 21-24), 

The origin of ray hairs is not so clear. In the Malayan-Asiatic 
line they seem to be derived from the basic type or one tending 
toward a glandular condition; there are all degrees of gradation 
between these types in P, grande and P. icandae {Figs. 27, 28). 
In the Afro-American line the ray hairs are often stouter, conical, 
and seem to have been derived from glandular hairs. These rays 
may appear as branches off a glandular hair in P. angolense, P. 
andimim {Figs. 14, 15), and occasionally in P. vassei, or may be 
independent but intermingled with glandular hairs in P. ellisii 
and P. vassei {Figs. 19, 20), Occasional hairs intermediate between 
the glandular and the ray condition may be found near the basal 
margin in P. vassei. The abundance of conical ray hairs also seems 
to indicate an intermediate condition betw^een glandular and ray 
hairs. When the contents of the conical ray hair collapse, the 
ordinary ray with its flat shape is produced. Conical ray hairs are 
mostly found in the Afro-American line, and basic hairs are mostly 
absent from it. In P. ellisii, the ray hairs tend to originate from 
the center of the trichoblast cell, rather than from one side, as is 
frequently the case with hairs close to the basic type in origin. 
Nayar (1965, p. 13) has described hairs on Pyrrosia obovata which 
seem to be similar in many ways to the ray hairs of Platycerium 
ellisii. Pyrrosia heteradis, with the basic type of hair, is considered 

Rhizome Scales of Platyceriuai 15," 

by Nayar primitive in relationship to Pyrrosia obovata. Nayar 
implied that the hairs which arise from the center of the cell 
and have dilated bases as in P. ohovata are separate entities not 
related to protuberances in origin. It seems that hairs of this type 
in Platijceriwn are a result of cell reduction or failure of the basal 
developing hair cell to elongate properly. Similar dihited hair 
bases on glandular hairs may be found in some forms of P, vassei 
and in P, ridleyi. 

Unicellular to multicellular hairs. — Most of the species have 
hairs with 2-4 cells. But one has only protuberances known as 
papillae (Fig, 25), others have mostly 1-celled hairs {Fig, 20) j and 
others have mostly multicellular hairs {Fig. 26). It is generally 
believed that unicellular hairs are more primitive than multicel- 
lular hairs unless the unicellular condition is shown to be a reduc- 
tion (Xetolitzky, cited in Carlquist, 1961, p. 34). In the [Malayan- 
Asiatic line of the genus the primitive P. ivallichii (Fig. 25) lacks 
hairs and has only papillae which presumably gave rise to 1-celled 
hairs. Hairs with more cells have evolved, and those of P. holttumii 
(Fig. 26) have 8 cells. The next species, P. waiidae and P. grande 
(Figs, 27 f 28), show a decrease in the number of cells to 5 or fewer. 
Basic hairs are still present, but semi-specialized hairs (i.e. inter- 
mediate glandular and ray types) have appeared. It seems that 
the appearance of semi-specialized to specialized hairs may be 
correlated with at least the partial loss of the basic hair types. In 
P. coronarium (Fig. 29) j an advanced species, the cells in the hair 
are reduced to 2 and are glandular. The more advanced P. ridleyi 
(Fig. SO) has a large percentage of 1-celled glandular hairs. This 
reduction trend is also found in the Afro-American line, culminat- 
ing in P, ellisii (Fig. 20) which bears only 1-eelled glandular or 



Incomplete stellate hairs. — Ray cells are unicellular (Fig. 19b) 




preaches the stellate hair. A less specialized development pro- 

156 AjNiEKiCAN Ferx Jourxal 

duces a long cell with many lateral ray-like cells along its axis; 
such hairs are occasionally found on fibrils. 

There seems to be a relationship between the stellate hairs 
found on the lamina and those on the rhizome scales. Rhizome 
scale hairs apparently show ontogenetic stages no longer present 
in the development of laminar stellate hairs. The development as 
studied by Straszewski (1915, p. 277) shows that a laminar stellate 
hair starts as a glandular cell w^hich divides apically into successive 
glandular cells that elongate and flatten into rays. By studying a 
variety of hairs on P. angolense scales it is possible to reconstruct 
a plausible seciuence of cell arrangement that could have led 
up to the stages described by Straszewski. 

One might take the point of view that the hairs on the rhizome 
scale represent reductions from the stellate condition. This seems 
unlikely, for the cells of a reduced hair would probably be com- 
prised of the component cells of the stellate hair type, i.e. ray 
and glandular cells. Instead, a continuous array of basic to special- 
ized types of cells are present. Also, if reduction were the case, 
the most primitive species would have retained more of the stellate 
hair structure. This is not the case, however, for the species havhig 
this condition are not among the very primitive ones. 

The stellate hair tendency in scales is best developed in P. 
angolense and P. andimim. The other species have other special- 
izations or have not reached the stellate state. The separation of 
the ray and glandular cells into independent hairs is one type of 
specialization and is found in P. ellisii {Fiy, 20), 

Flabelloid niargin.~~The cells of the scale margins are generally 
arranged longitudinally, but may be oriented at various angles, 
and may even be retrorse (Figs, 2, 28). Margins with cells that 
spread outwards in a fan-like manner Holttum (1057, p. 43) 
described as flabelloid. Flabelloidly arranged cells may be absent, 
in clusters, or continuous, well developed or barely so, and they 
may merge with papillae and trichoblasts. 

If cells bearing protuberances, trichoblasts, and basic hairs are 
essentially similar and if they can unite laterally to each other in 
varying degrees, it is possible to explain the origin of the flabelloid 

Rhizome Scales of PLATYCERimi 157 

margin by lateral fusion of "free" marginal cells. This hypothesis 
is likely because there is no difference in color and texture between 
"free" marginal cells and those making up the flabelloid margin, 
the marginal cells being united in varying degrees up to a typical 
flabelloid state, and the orientation of the cells in the flabelloid 
margin being in the same direction as that of **free" hairs. That 
marginal cells as well as submarginal cells of a flabelloid margin 
may continue to produce papillae and hairs is further evidence 
that the cells of the flabelloid margin are basically the same as the 
marginal cells of unmodified scales. If the ''free" hairs on a flabel- 
loid margin unite, a very broad, flabelloid margin as found in P, 
coronarium and P. ridleyi is formed (Fig. 2), The cells of these 
broad, flabelloid margins are oriented like many of the ''free" 
marginal hairs in P. grande. In sunmiary, the trend in the Malayan- 
Asiatic line seems to be from marginal cells arranged longitudi- 
nallj" to a narrow flabelloid margin with many "free" hairs to a 
wider flabelloid margin with few "free" hairs. This trend is absent 
or poorly developed in the other evolutionary lines of the genus. 
Where it does appear it is found in advanced species (P. vassei 
from ^fadagascar, P. bifurcaturrtj and P. hiUii). 

False branching. — False branching occurs in P. holttumii (Fig. 
26) and occasionally in P. grande. It is characterized by having 
one or more hair cells with protuberances. Occasionally the pro- 
tuberances are separated from the cell by a wall, and then a 
truly branched condition exists. False branching may be inter- 
preted as a stage toward the fusion of hair cells to form a flabelloid 
margin. An alternative explanation is that these relatively newly 
derived basic hairs are still genetically close to the marginal cells 
and have retained their tendency to form papillae or, alternately, 
false branches. This seems to be the more adequate explanation, 
as many of the false branches are not distinct from the papillae 
of marginal cells. 

Function of the Rhizome Scales 

The most vulnerable part of these sometimes immense plants is 
the relatively small apical bud. Death of the bud results in death 

158 American Fern Journal 

of the whole plant in species which do not produce lateral buds or 
buds from the roots. Rhizome scales function to protect the bud 
from desiccatioji, animals, and excessive water. Beneath the base 
fronds old rhizome scales may be found very much intact, covering 
and protecting the rhizome, although the softer parts of the scales 
may be worn away leaving only the sclerified stripes. 

Protection from desiccation is particularly important in these 
usually high epiphytes, as they are exposed to more drying condi- 
tions than low epiphytes or terrestrial plants. I do not know what 
animals might attack these plants in their native habitats, but the 
mass of base fronds is known to be a home for a multitude of 
organisms. Sowbugs, pillbugs, snails, and slugs are capable of 
eating through the scales and into the buds, particularly on young 
plants in cultivation. Small insects would probably have difficulty 
breaking through the scales covering the bud. I have observed 
scale insects unable to feed because they were entangled in the hairs 
of the rhizome scales. The insects were able to attach and feed 
where the rhizome scales ended on the phyllopodia. Smaller 
animals, however, could gain entry to the bud and rhizome through 
the injury inflicted by larger animals or micr organisms. The small 
larvae of fungus gnats {Mycelophilidae) apparently gain entry to 
the rhizome bud through leaf scars which are perhaps softened by 
decay or do not suberize well. Termites, birds, and various fungi 
also have been observed to make their home in the nest fronds as 




water. I believe that keeping excess water off the bud tissue is a 
very important function of the rhizome scales as well. Water- 
logged tissues are particularly vulnerable to microrganism infec- 
tions. If softened areas are produced from such infections, other 
organisms then readily find their way into the remainhig tissue. 
The glandular hairs and large amounts of oil found in the rhizome 
scales of P. hifurcatum^ P. grande (Joe, 1964, p. 71), and probably 
other species must also play some role in rhizome protection. 

Rhizome Scales of Platyceriuai 159 


With a knowledge of the rhizome scale characters described 
above and thek distribution among the 17 species, it is possible 
to infer the kind of scales most likely possessed by the ancestral 
plant. In Table I, P. wallichii has the lowest value for specialized 
rhizome scale characters and probably has scales most closely 
resembling those of the ancestral plant. The relationships of 
present day species also may be clarified. For example, the general 
appearance of the South American species P. andinum could place 
it in any of the three evolutionary lines of the genus, but several 
rhizome scale characters and other less obvious structures place 
it closest to the African species. However, of broader interest are 
the changes in rhizome scale structures that have taken place along 
with the general evolution of the various species. Some of these 
changes are limited to a few species, but others are established 
patterns in this and probably other fern genera. 

The following changes which have taken place in rhizome scales 
of Platycerium seem to be correlated with evolution in the genus: 
fibrils present to absent; width narrow to broad; texture firm to 
thin papery, at least as a broad margin; apex fiUform to non-fih- 
form and multicellular; base broadly attached to subpeltately to 
peltately attached; hairs restricted to the scale margin to spreading 
submarginally or centrally; margins mostly with papillae to mostly 
with basic hau-s to mostly with typical rays and/or glandular 
hairs; hairs absent or basic hairs mostly 1-celled to basic hairs 
multicellular; simple glandular or ray hairs with many cells to the 
same with few cells; simple hairs to partly formed stellate hairs; 
and marginal cells oriented with the length of the scale to flabel- 

loid to very broad flabelloid. 
From the rhizome scale characters having phylogenetic value 

the following generahzations are made: 

1. Elaborate scale structure is indicative of advanced species, 
except where reduction of the hair cells is involved.' 

' A .similar conrlusion is reached by R. M. Tryon, Jr., who finds th^i Cyathea 
scales with the least cellular differentiation are primitive (m Ittt. &ept. », 
1969) . 


160 American Fern Journal 

2. Except in very primitive species, reduction in number of hair 
cells of simple but not stellate hairs is apparently correlated with 
increased hair specialization. 

3. Hair development on some rhizome scales seems to reflect 
stellate hair ontogeny and shows stages now absent from the leaf. 

4. The flabelloid margin seems to have developed from the union 

of papillae, trichoblast, and hairs. 

5. Peltate scales developed by the folding under of the auriculate 
or rounded base and subsequent overtopping growth, rather than 
by the lateral union of the auricles. This is similar to the formation 
of peltate scales in Pyrrosia, 

6. One of the important functions of the rhizome scale may be 
to protect the bud tissue from becoming water-logged. 

It seems that greater use may be made of rhizome scales in 
determining relationships within other genera, providing sufficient 
features are available. Scale interpretations will be more reliable 
with some prior knowledge of at least some of the relationships 
in the group. The use of scale characters will be of greatest help 
in interpretation of problematic taxa and in reinforcing or support- 
ing other phylogenetic interpretations. 

Literature Cited 

Bower, F. O. 1923. The Ferns, vol. I. 359 pp. University Press, Cambridge, 
Carlquist, S. 1961. Comparative Plant Anatomy. 146 pp. Holt, Rinehart, 

and Winston, New York. 


Joe, Barbara. 1964. A Review of the Species of Platycerium (Poly podiaceae) . 

Baileya 12: 69-126. 
Nayar, B. K. 1961. Studies in Polypodiaceae, VII: Pyrrosia. J. Indian Bot. 

Soc. 40: 164-186. 
Nayar, B. K. and S. Chandra. 1965. Ferns of India-XV, Pyrrosia. Bull. 

Nat. Bot. Card., Lucknow 117: 1-98. 
Straszewskt, H. R. 1915. Die Farngattung Platycerium. Flora 108: 271-310 

Wagner, W. H., Jr. 1952. The Fern Genus Diellia, its Structure, Affinities, 

and Taxonomy. 167 pp. Univ. Calif. Publ. Bot. 26(1): 1-212. 
Wagner, W. H., Jr. 1964. The Evolutionary Pattern of Living Fern: 

Torrey Bot. Club 21 : 86-95. 

Los Angeles City College, Los Angeles, California 90029. 

Shorter Notes 101 

Shorter Notes 

Stem Cross-section Prints Aid in Identifying Horsetails, 
— Stem cross-sections are often required for accurate identification 
of horsetails. These are difficult to obtain from dry and brittle 
herbarium specimens. I have found a method for making a printed 
record of the cross-section of a horsetail specimen (luickly and with 
simple materials. It must be done before the specimen is pressed, 
but maj^ be affixed to the herbarium sheet later as an accurate 



is ready to go into the plant press, a sharp, transverse cut is made 
across the center of a mature internode. One of the cut surfaces 
is inked with a^ fountain or felt pen. Several imprints of the surface 
can then be made on smooth, non-absorbant paper (the field label 
if possible). Rough paper will make details difficult or impossible 
to see, and absorbent paper will change the print's dimensions. 
Although the first prints may be inked too heavily, others will be 
perfect, and some Avill be too light. Unless a good labelling system 
is used to associate the prints with the particular specimens, 
the prints should be kept with the specimens in the press. The 
best prints should be pasted near the specimen label when the 
specimen itself is mounted. — Barbara J. Gudmundson, 5505 28th 
Avenue South, Minneapolis, Minn. 55417. 

Psilotum nudum Spreading Northward. — In 1942 the ''Whisk 
Fern/' Psilotum nudum, was recorded in Brown and Correll's 
'Terns and Fern Allies of Louisiana'' as having been collected 
growing wild in a greenhouse, Ivrack*s Nursery, north of New 
Orleans, having apparently been accidentally imported with 
potting soil. A recent publication (Sida 3: 525. 1970) by Dr. G. 
Rhodes now reports the species as growing ^ild in four Louisiana 
Parishes^Terrebonne, St. INIary, Vermilion, and Lincoln. In the 
last named the species is some 250 miles north of any previously 
known locaUty. It is thus apparent that this plant is spreading 


eastw^ard in Alabama or Mississippi.^C.V.^I. 

1C2 American Fern Journal 

Terminology of the Sporanglvl Structures of Equisetu:vl 
Several years ago a population of Equisetum X Utorale with 
many abnormal cones was discovered by Dr. W. H. Wagner, Jr., 
near Milan, Michigan, The population was extensive, and had 
invaded an old bean field. In this dry, sandy habitat, atypical for 
this hybrid, the strobiliferous stems frequently had cones with 
proliferated tips {Fig. 1), The apex of the cone, instead of ceasing 
growth as it usually does, reverted to a vegetative condition. This 
type of teratology has been previously recorded in various species 
of Equisetum, and Gluck,^ Kashyap,^ and Tschudy^ have described 
the intergradation of fertile and sterile appendages associated with 
the transition from the strobiUferous to the vegetative apex. The 
same intergradation, seen here in E, X Utorale {Figs, ^-7), reveals 
an innate capability of the appendages produced by the apex to 
develop either as vegetative leaves or as sporangium-bearing 
structures. This indicates that the two structures are homologous, 
and that the term '"sporophyll," a more exact term than "spo- 
rangiophore," should be used for these structures. Tschudy argued 
for the term ^^sporophyll" on the basis of teratology in E, telniateia, 
and I wish to reiterate it here on the basis of teratology in E, X 

Since the leaves of Lycophytina, Sphenophytina, and Pterophy- 
tina are all apparently independently evolved, calling their 
sporangium-bearing structures ''sporophylls" does not imply any 
homology between the sporophylls of these groups, but only be- 
tween the leaves and sporangium-bearing structures within each 
group. — Richard L. Hauhe, Department of Botany, University 
OF Rhode Island, Kingston, R. I. 02S8L 

> Die Sporophyllmetamorphose. Flora 80: 303-387. 1895. 
* Some abnormal cones in Equisetum debile. J. Indian Bot. Soc. 9 : 240-241. 

^ The significance of certain abnormalities in Equisetum. Amer, J. Bot. 26: 
744-749. 1939. 

American Fern Journal 

^\)LUME 60, Plate 20 


2 cm 

J i 

Fig. 1. Cones of E. X litorale showing proliferation of the tips. 
Figs, 2-7. Series of selected appendages from the cones in Fig. 1, from 
A typical sporophyll to a nearly typical sheath segment (leaf). 

164 American Fern Journal 

Kinds of Cystopteris in California. — Recently John Thomas 
Howell, while preparmg a floristic account of the ferns of the 
Sierra Nevada, suggested to me a study of the two spore types of 
Brittle Bladder Fern in that area. I enlarged the project's scope 
to include all the Cahfornia collections of Cystopteris in the Cali- 
fornia Academy of Sciences herbarium, more than 120 spore- 
bearing collections. 

Based on spore characteristics, two separate taxa are recogniz- 
able, although in general appearance they look identical. Profumo^ 
examined the gametophytes of the tw^o types and also found no 
morphological differences. At times, the two types have been 
considered separate species, C. fragilis (L,) Bernh., wdth spiny 
spores, and C dickieana Sim, with non-spiny (rugose-verrucose) 
spores; the two have also been considered subspecies. 

The specimens I examined show that both kinds may occur in 
the same locality, a fact which was recognized byProfumoand by 
Hagenah,^ w^ho occasionally found both mounted on the same 
herbarium sheet. 

Within the Sierra Nevada, 14 collections were the fragilis type 
and 67 the dickieana type, a noteworthy excess of the latter, Non- 
Sierran collections showed 22 fragilis type and 20 dickieana type, 
a more even distribution. While both kinds occasionally may occur 
together, there appears to be a predominance of the dickieana 
type at higher elevations. This corresponds to the findings of 
Larsen^ in Greenland, w^ho suggested that C. dickieana w^as a high 
Arctic species and that C fragilis was a sub- Arctic species. 

Perhaps these two taxa should be considered as forms rather 
than subspecies since their distributions are essentially identical 
and the only apparent difference is in their spores. — Robert B. 
Setzer, Allan Hancock Foundation, Dept. of Biology, Univ, of 
Southern California^ Los Angeles, California 90007. 

^ — ^ 

^ Prof umo, P. 1969. The gametophj^te of Cystopteris fragilis and Cystopteris 
dickieana. Webbia 23 : 317-328. 

*Hageiiah, D. J. 1961. Spore studies in the genus Cystopteris. I. The distri- 
bution of Cystopteris with non-spiny sp6res in North America. Rhodora 63: 

2 Larsen, K. 1952. Udbredelsen i Gronland af Cystopteris fragilis coll. med 
piggede og vortede sporer. Bot. Tidsskr. 49; 39-44. 

Notes and News 165 

Notes and News 

A New Newsletter. — To better promote comminiication and 
the exchange of materials between our members, the American 
Fern Society is inaugurating a quarterly Newsletter. Prof. James 
Montgomery, an enthusiastic student of North American fenis 
and a frequent participant in our Fern Forays, and ]Mr. F. Gordon 
Foster, an experienced horticulturist and author of ^*The Garden- 
er's Fern Book," will be in charge of the Newsletter. Non-technical 
items and notes of passing interest will be most welcome. The 
officer's reports and a membership list will appear in the News- 
letter each year. We hope especially that our members who are 
naturalists and gardeners will contribute to the Newsletter and 
will find it interestino; and informative. 

George Neville Joxes, Professor of Botany and Curator of 
the Herbarium at the University of Illinois, died last June 25. 

LB ^^ 

He had been a Fern Society member for a quarter of a century. 
He was the author of many monographs and floras, including "A 
Botanical Survey of the Olympic Peninsula" and '''Flora of Illi- 
nois." His principal work on ferns, "An Annotated BibHography 
of ]Mexican Ferns," was published four years ago. 

Plaxts of Asplexium ebenoides X Camptosorus rhizophyl- 
Lus are available to persons interested in growing this backcross 
hybrid Spleenwort. Please request living specimens. Scott's 
Spleenwort, A . X ebenoides, is the hybrid of the Ebony Spleenwort, 
A. platyneuron, and the Walking Fern, Camptosorus. We have 
available numerous prohferations of the backcross of the hybrid to 
the walking-fern, which was originally made by Airs. Kathryn E. 
Boydston at Fernwood, Niles, Michigan. This backcross plant has 
proved to be extremely vigorous in cultui'e. It looks like a some- 



or terrarium in ordinary potting soil. — W. H. Wagxer, Jr., 



166 American Fern Journal 

Edward M. Shields, 1895-1970. — Alembers of the American 
Fern Society, especially those who some years ago participated in 
its field trip in eastern New York, will regret to learn of the death 
from a heart attack on September 17th of * 'Eddie'' Shields, who 
on the occasion of the field trip entertained us at the summer 
residence he then maintained at Hames Falls in the Catskills. 

As a youth Eddie roamed his native Chester County, Pennsyl- 
vania, collecting at first minerals and Indian relics. Receiving 
from the family real estate holdings there sufficient income to live 
comfortably, he became an amateur naturalist of wide interests. 
On a field trip in Florida he made some notable orchid finds, 
thereby becoming acquainted with Oakes Ames, the specialist on 
these plants. His interest in ferns was aroused when, on a visit 
to Cambridge, Ames introduced him to Charles A. Weatherby. 
Exploring the Haines Falls region for ferns, he discovered a remark- 
able southern disjunct outlier of the Fragrant Fern, Dryopteris 
frag vans, 

^ w 

After some years, Eddie came to spend his whole time in Sara- 
sota, Florida. Studying subtropical horticulture, he developed 
around his house a garden including many uncommonly used yet 
highly attractive plants. In an adjoining slat-house, which after a 
couple of disastrous freezes evolved into a greenhouse, he special- 
ized on growing unusual orchids. He would stop in at one of the 
many Florida orchid-sales places, and with his keen eye and thor- 
ough acquaintance with current literature, would often detect as 
a rarity a plant offered as a common one. 

Ferns in his garden greenhouse were merely incidental; in the 
living room he had a spectacular hanging-basket culture of Narrow 
Strap-fern (Campyloneuron angustijolium) . His fern books and 




and stepchildren. Memories of his cordiality and generosity to me 
over the years have made this note most difficult for me to write. 

19 10 A 

)f Penr 

Recent Fern Literature 167 

Recent Fern Literature 

Pacific Northwest Ferns and Their Allies, by T. M. C. 
Taylor. University of Toronto Press, 33 East Tiipper St., Buffalo, 
N. Y, 14203. 1970. 247 pp., illustr. Slo.OO.— This is the first full- 
fledged local fern flora to appear in several years. The area covered 
includes that treated recently in the '^Vascular Plants of the 
Pacific Northwest'' (reviewed in this Journal. 60: 34-38. 1970), 
but is more extensive, since all of British Columbia, the Yukon 
Territory, and all of Alaska is included. The number of species 
treated is 97, about a quarter of all those known from North 
America north of Alexico. ]\Iany of these are boreal plants familiar 
in the eastern United States and Canada and manj^ occur also in 
Eurasia. Twenty-seven are western American endemics (but not 
local endemics, except for Botrychium piimicolay confined to the 
region of Crater Lake, Oregon). The species all have full descrip- 
tions and most of them have distribution maps and line drawings. 
The latter are very good, so good that most species can be easily 
recognized from the drawings alone. (I would except the drawing 
of Polystichum scopulinum, which shows a plant not typical of this 
species and which might even represent P, kruckebergii instead.) 
The systematic treatment is commendably conserv^ative. The 
comments indicate the variations found and some of the taxonomic 
problems still unsolved. There are few errors indeed, and the 
comments below are of strictly minor importance. 

The authority of Blechnum spicant is not ''J. Sm.'' but Roth 
(cf. this Journal 34; 51. 1944); hicidentally, it is usual to use "J. 
Sm." for John Smith; the author intended here was not John 
Smith but Sir James Edward Smith, usually cited as J. E. Smith, 
for the abbreviation of Smith to '*Sm." is really not sanctioned by 
the Code. The work in which Dryopteris was published by Adanson 
is not 'Tarn PL" but 'Tam. PL" In the description, the sori of 
Polypodium montense are said to be ''submarginal," but in the 
drawing they are shown as about medial (for a comment on the 
name "montense'' see this Journal 60: 126-7. 1970). Polypcdium 
virginianum is described as having the segments ''linear-oblong to 

168 American Fern Journal 

lanceolate . . . obtuse at apex/' which is essentially correct, al- 
though the apex does vary from rounded to acute, but in the com- 
ments it is said to be distinguishable from P. montense by the 
segments being ''narrowly ovate with acute tips/' which is not true 
and which contradicts not only the description but also the draw- 
ingj which shows the segments as oblong and rounded at apex. In 
the synonymy of Thelyptens nevadensis, the first line ought really 
to be ''ex ]\Torton, Amer. Fern J. 48: 139. 1958/' as required by 
the Code; otherwise it appears that Clute published the name 
some eight years after his death. The Azolla of the area is referred 
to A. mexicana, following Svenson; the report of A. filiculoides 
from Alaska by Svenson is rejected with the statement ''almost 
certainly an ac^uarium escape/' which is perhaps an unwarranted 
assumption, since the specimen was collected in 1868, an early 
period in the history of Alaska when aquaria must have been 
essentially unknown; still, the record must be considered doubtful 
since no other Alaskan material has ever turned up. There are 
specimens in the U. S. National Herbarium from Oregon and 
Washington that have been identified as A . filiculoides (and sub- 
sequently to Svenson's treatment), but they are sterile, I believe, 
and so somewhat doubtful. An effort should be made to collect 
fertile material from these states. Possibly Svenson overstressed 
the character of the septae in the glochidia, and his treatment has 
never been critically evaluated. Although it is irrelevant here, I 
might mention that although Svenson indicated that -4. caroliniana 
must spread solely by vegetative means, since megaspores are 
never found in American material, some recent European workers 
have described the megaspores of this species, considered intro- 
duced in Europe. It is possible that the European plants so identi- 
fied are not really A^ caroliniana^ and the matter should be in- 
vestigated. It is rather too bad that this book perpetuates the 
spellings "sa6i/m^/ohwm," ^^matricariaefolium,^^ and ^^andromedae- 
folia''; although these were the original spellings, Art. 73, Note 2, 
says that the use of a wrong connecting vowel is to be treated as 
an orthographic error, and Art. 73 itself indicates that orthographic 
errors are to be corrected; thus the correct spellings are '^sahini- 

American Fern Society 169 

foliurrij^^ ''matricarnfolmm/' and ^'andromedifolia.^^ One feature, 
which does not at all detract from the book but which is sufficiently 
peculiar to deserve mention, is the placing of SelagineUa at the 
end, following the Polypodiaceae, instead of at the beginning, 
with Lycopodium, Isoetes, Equisetum, and other fern allies. — C.V.^I. 

American Fern Society 

New Members 

Mrs. Rhoda D. Bancroft, 4 Willard Ct., Norwich, NY 13815 

Mrs. Henry W. Bookout, Jr., 120 Smith St., Islip, NY 117.51 

Air. Parul Catling, 104 Victoria Park Ave., Toronto 13, Ont., Canada 

Mrs. Thomas A. Cole, 1000 S. Grant Ave., Crawfordsville, IN 47933 

Mr. Edward L. Davis, Dept. of Botany, Univ. of Massachusetts, Amherst 

MA 01002 
Mr. Robert B. Faden, East African Herbarium, P. O. Box 5166, Nairobi, 

Kenya, Africa 
Mrs. Leslie J. Helmstetter, 2546 Hillsman St., Falls Church, VA 22043 
Mrs. J. J. Howell, 4028 Greenwood Dr., Ft. Pierce, FL 37450 
iMrs. S. H. Inglis, 233 Red Fox Road, Stamford, CT 06903 
Mr. R. Kean Ivey, Jr., Box lOS, Rt. 3, Charlottesville, VA 22901 
Mr. Terry W. Lucansky, 3600 Tremont Dr., Apt. H-3, Durham, NC 27705 
Mr. Donald IMcFall, 4711 Forest Hills Rd., Rockford, IL GUll 
Mrs. Robert D. Moody, 7705 Lynch Rd., Sebastupol, CA 95472 
Mr. Richard Moyroud, 202 Grove Way, Delray Beach, FL 33444 
Mrs. Jane W. Pettus, S05 South Warson Rd., St. Louis, MO 63124 
Mr. Michael G. Price, Dept. of Botany, UPCA, College, Laguna, Philippines 
Mr. Arthur Raguse, 715 Linwood St., Abington, MA 02351 
Rev. Paul John Rich, First Parish Unitarian Church, E. Bridgewater, MA 


Mrs. William J. Wright, 1914 W. Division St., Arlington, TX 76010 

Changes of Address 

Mr. Michael I. Cousens, Dept. of Botany, Washington State Univ., Pullman, 

W A 99 1 63 

Mrs. V. De Benedictus, Zoology Dept., Syracuse University, Syracuse, NY 


Dr. Delzie Demaree, 109 South Ave., Hot Springs, AR 71901 

Mr. William E. Fulcher, Biology Dept., Guilford College, Greesnboro, NC 



American Fern Journal 

Index to Volume 60 

Abacopteris, 120 

Aberrant Leaves on Angle-Shoots of Sela^ii- 

nella martensii Spring, 1 
Actiniopteris radiata, 137, 138 
Acrostichum alcicorne, 7-11; bifurcatum, 9; 

stemaria, 7, 9, 11 

Adiantaceae, 123 

Adiantum, 124, 137, 138; aethiopicum, 138j 
capillus-veneris, 128, 137, 138; caudatum, 
137, 138; lunulatum, 137, 138; pedatum, 
33; vennstum, 137, 138 

Aleuritopteris, 125 

Allen, D, E. The Victorian Fern Craze 

(rev,), 84 

Anogramma leptophylla, 101-103, 128 

Anopteris hexagona, 103 

Apalophlebia costata, 121 

Aspidiaceae, 34, 73 

Aspidium, 34; spinulosum, 24; thelypteris, 24 

Aspidotis, 125; densa, 40 

Aspleniaceae, 35 

Asplenium, 86, 124; adiantum-nigrum, 138 
subsp. onopteris, 128; X germanicum, 133 
falcatum, 139; incisum, 86; indicum, 121 
longissimum, 121; oligophlebium, 86 
platynenron, 131, 133; ruta-muraria, 139 
septentrionale, 129-133; sylvaticum. 122 
trichoraanea. 131, 139; tripteropus, 86 
varians, 103; vestitum, 122 

Athyrium, 34, 38; filix-femina, 139, cv. 
'Corymbiferiim,' 28; japonicum, 100; 
lilloi, 103; thelypterioides, 100 

AzoUa, 35; caroUniana, 168; fiUculoides, 168; 
mexicana, 168 

Azollaceae, 123 

Bajpai, Nisha (see B. K. Nayar) 

Bslknap, Frances (see L. Lee) 

Bibliography to Floras of Southeast Asia 
(rew), 39 

Bir, S. S. A Note on the Gametophytes of 

Cystopteris tennisecta, 98 
Blechnopsis malaccensis. 121 
Blechnum, 104; andicola, 105; araueanum, 

105; australe, 104; doodioides, 36; gaya- 

num, 104;gerniainii, 104, 105; indiuum, 121; 

tnagellanicum var. angustiseta, 105, van 

setigerum. 105; microphyllum, 103, 104; 

nivale, 105; penna -marina, 103, 104; 

poeppigianuin, 104; serrulatum, 121; 

spicant, 35, 167; tabulare var. setigerum, 


Botrychium. 62. 63, 139; lunaria. 139; 

pumicola, 167; ternatum, 139; virginianum, 

62-64. 139, var. gracile, 62 
Camptosorus, 126 
CardiUo, Frances M. Stelar Anatomy of 

Six Species of Lycopodium, 89 
Ceterach officinarum, 128 
Chandra, S. Vascular Organization of the 

Rhizome of Cibotium barometz, 68 

Cheilanthes, 125; catanensis. 128; fragrans, 

128; gracillima, 40; lanosa, 36; siliquosa, 

Chromosomes and Speciation in Ferns (rev.), 

Cibotium, 68; barometz. 68. 70, 71. 139; 

subg. Microcibotium. 68 
A Commentary on Some Type Specimens of 

Ferns in the Herbarium of K. B. Presl 

(rev.), 119 

Cornopteris, 98, 100 

Cousens, M. I. & H, T. Horner. Gametophyte 
Ontogeny and Sex Expression in Dryo- 
pteris ludoviciana, 13 

Cronriuist, A. (see C. L. Hitchcock) 

Cryptogramma, 60 

Ctenopteris, 123 

Cyathea, 38, 71, 159 

Cyatheaceae, 68, 71 

Cyclophonis princeps, 118 

Cyclosorus, 126; megaphyllus, 120 

Cystopteris, 98. 100, 164; subg, Acystopteris, 
98, 100; bulbifera, 98; subg. Cystopteris, 
98; diaphana, 98, 100; dickieana. 164; 
douglassii, 98, 100; fragilis. 98, 100. 164; 
montana, 35; tenuisecta, 98-100 

Dennstaedtia, 33, 39 

DeVoi, C. E. The Pteridophytaof Tai\van-5. 
Hymenophyllaceae: Trichomanes (rev.), 87 

Dicranopteris, 38 

Didymoglossum magellanicum, 106 

Diellia, 144 

Diplaziopsis javanica, 100 

Diplazium, 38; esculentum, 98;grammitoides, 
122; sylvaticum, 122; tenerum, 122; 
vestitum, 122 

Dipteris, 125; conjugata, 125 

Drynaria quereifolia, 120, 139 

Dryopteridaceae, 35, 116, 117 

Dryopteris, 14, 24, 167; assimilis. 38; aus- 
triaca, 37, var. austriaca, 37; barbigera, 
137; carthusiana, 33, 38; chrysocoma, 137; 
cristata, 33; dilatata, 38; filix-mas, 24, 25, 
137. 139, 140; intermedia, 33; ludo- 
viciana. 13-21, 23-26; marginata, 137; 
odontoloma, 137; ramosa, 137; schim- 
periana, 137; spinulosa, 24. 33, 38; viilaru 

subsp. australis, 128 
Elaphoglossum, 73, 74, 77-81, 125; auripilum, 

77; conspersum, 75; denudatum, 103; 

erinaceum, 75; eximium, 75; herminieri, 75; 

hirtum, 75; hyatinum, 75; lindenii, 75; 

longifolium, 75; muscosum, 75, 77; revol- 

vens, 75, 77; squamipes, 75; tectum, 75, 77; 

undulatum, 79 
Elmore, H. W. Two New State Records for 

Isoetes in Kentucky, 30 

Emory, D. L. A Major North American 
Range Extension for the Forked Spleen- 
wort, Asplenium septentrionale, 129 

Equisetum, 162, 169; ar\^ense, 137, 140; 
debile, 140; X litorale, 162, 163; ramosis^- 
simum, 128; telmateia, 128, 162 

The Fern Collections in Some European 

Herbaria, VI, 49 
Fern Hybridizing at the University of Leeds 

(rev.), 86 ... 

Fertile Sporophytes of Botrychium vir- 
ginianum Attached to Gametophytes, 62 

Flora Palaestina, Part One, Text (rev.). 127 

A Further Note on the Type of Platycenum 
alcicorne, 7 

Gametophyte Ontogeny and Sex Expression 

in Drypoteris ludoviciana, 13 
Gleichenia, 38 

Goniophlebium, 66; villeminianum, 67 
Goniopteris tetragona, 120 
Grammitidaceae, 123 
Grammitib, 123; albidula, 66; crispata, 66; 

sect. Cryptosorus, 66; discolor, 66; eminens, 

Index to Volume 60 


66; kalbreyeri, 66; mathewsii, 66, 67; 
sechellarum, 124; trifurcata, 66 
Gudraundaon, Barbara J. Stem Cross- 
section Prints Aid in Identifying Horse- 
tails. 161 

Hard Water as a Limiting Factor in the 
Distribution of Isoetes echinospora, 134 

Harrison. S, G. (see H. A, Hvde) 

Hauke, R. L. Terminology of the Sporangial 
Structures of Equisetum, 162 

Helminthostachys zeylanica, 140 

Hitchcock, C. L., A. Cronquist, M. Ownbey, 
& J. W. Thompson. Vascular Plants of the 
Pacific Northwest, Part 1 (rev.), 34 

Holttum, R. E. A Commentary on Some 
Type Specimens of Ferns in the Her- 
barium of K. B. Presl (rev.), 119; A Revised 
Flora of Malaya, vol. II— Ferns of Malaya, 
ed. 2 (rev)., 38 

Horner, H. T. (see M. I. Cousens) 
Hoshizaki, Barbara Joe. The Rhizome Scales 

of Platycerium, 144 
Hyde. H. A. & A. E. Wade. Welsh Fern^, 

Clubmosses, Quilhvorts, and Horsetails, 

A Descriptive Handbook, ed. 5 (rev.), 84 
Hymenophyllum, 106; magellanicum, 105, 

106; seselifolium. 105 
Hypodematium, 107, 114-117; crenatum, 98, 

100, 107. 108. 112, 115 
The Illustrated Flora of Illinois: Ferna (rev.), 

Indian Pteridophytes Used in Folk Remedies, 

Isoetes, 30, 35, 169; sect. Echinatae, 37; 

echmospora, 37, 134, 135, var. braunii, 37. 

135, 1 36 ; engelmannii, 30 ; melanopoda, 

30; muricata, 37; setacea, 37; sect. Tuber- 

^culatae, 37; tenella, 37 

Karyological Studies on Aspleniaceae 1. 

Karyotypes of Tliree Species in Asplenium 
(rev.), 86 

Kawakami, S. (see S. Tatuno) 

Kinds of Cystopteris in California, 164 

Lang, F. A. A New Name for a Species of 

Polypodium from Northwestern North 

America (rev.), 126 
Lastreopsis, 115 

The Lectotj-pe of Polypodium leptophyllum 
L., 101 

Lee, L. &. Frances Belknap. Hard Water as a 
Limiting Factor in the Distribution of 
Isoetes ecltinospora, 134 

Lindsaea, 124 

Lloyd, R. M. A Survey of Some Morpholo- 
gical Features of the Genus Elaphoglossum 
m Costa Rica, 73 

Lomariopsidaceae, 123 

Lomaria australis, 104; gayana, 104; ger- 
mainii, 104, 105; magellanica var. angua- 
tiseta, 105; microphylla, 103, 104; setigera. 

Lovis, J. D. Fern Hybridizing at the Uni- 
versity of Leeds (rev.), 86 

Lycopodium, 35, 89, 91, 93, 97, 141, 169; 
annotinum, 90, 141, var. annotinum, 91, 
95; cernuum, 140; clavatum, 91, 95, 96. 
137, 140; complanatum, 141; flabelliforme, 
91, 94, 95; lucidulum, 90, 91, 95, var. 
tryonii, 33; obscurum, 91, 92, 95; poro- 
philum, 33, 131; selago. 141; tristachyum, 
91, 95 

Lygodium, 38; circinnatum, 141; flexuosum, 

141; japonicum, 141; microphyllum. 141 
A Major North American Range Extension 

for the Forked Spleenwort, Asplenium 

septentrionale, 129 
Marsilea, 35; minuta, 128; quadrifolia. 141 
Mertensia magellanica, 106; quadripartita, 


Microgramma, 125 

Microlepia, 28; speluncae, 28, cv. •Corymbi- 
fera*, 28, 29; strigosa, 28, cv. 'Cristata.' 28 
JMicrosurium, 125, 126 

Mitui, K. Chromosomes and Speciation in 
Ferns (rev.), 39 

Mohlenbrock, R. H. The Illustrated Flora 
of Illinois: Ferns (rev.), 32 

Morton, C. V. A Further Note on the Type 
of Platycerium alcicorne, 7; The Lectotype 
of Polypodium leptophyllum L., 101; A 
New Form of Microlepia speluncae, 28; A 
Peculiar Species of Grammitis, 65; Psilo- 
tum nudum Spreading Northward, 161; 
Pyrrosia princeps, a Fern New to Culti- 
vation, 118; Taxonomic Notes on Ferns, 
IV, 103 

Nayar, B. K. A Nisha Bajpai. A Rein- 
vestigation of the Morphology of Hypo- 
dematium crenatum, 107 

Nephrolepis, 116 

Neuroplatyceron, 7 

A New Form of Microlepia speluncae, 28 

A New Nauie for a Species of Polypodium 
from Northwestern North America Crev.), 

Niphobolus costatus, 121 

A Note on the Gametophytes of Cystopteris 
tenuisecta, 98 

Notholaena, 125; vellea, 128 

Ophioglossum, 38, 60; lusitanicum, 128; 
vulgatum, 141 

Oleandra, 81 

Osmunda regalis, 141 

Ownbey, M. (see C. L. Hitchcock) 

Pacific Northwest Ferns and Their Allies 
(rev.), 167 

A Peculiar Species of Grammitis, Qo 

Pellaea, 60 

Phegopteris connectilis, 131 

Phillips, W. S. Psilotum nudum New to 
Arizona, 30 

Phyllitis sagittata, 128 

Phymatodes, 125, 126; banerjiana, 120; 

nigrescens, 120 
Platycerium, 121, 144, 155, 159; aethiop- 

icum, 11, 12; alcicorne, 11; andinum, 141- 

146, 148, 152, 154, 156. 159; angolense, 11. 

12. 144-146, 148, 150-152. 154, 156; bi- 

furcatum. 11, 146, 148, 149, 153, 157. 158; 

coronarium. 146, 147, 150, 153, 155. 157; 

elephantotis. 12; ellisii, 144. 151. 153-156; 

grande. 146, 147, 149-151, 153-155, 157. 

158; hillii, 146, 148^150, 152. 153. 157; 

holttumii, 146, 150. 151, 153-155, 157; 

madagascariense, 146; quadridichotomum. 

144, 146, 152; ridlevi, 146, 147. 151, 153, 

155, 157; stemaria, 11. 12, 146, 152; vassei, 

9, 11, 12. 146, 151-155, 157: veitchii, 146, 

148, 150, 153; wallichii, 146, 149. 153, 155. 

159; wandae, 146, 147, 149-155; wilhn- 

ckii, 146-148, 152, 153 
Pyrrosia, 149, 151, 160; heteractis, 154; 

obovata, 1.54, 155 


American Fern Journal 

Pleopeltis, 125 
Polypodiaceae, 34, 106, 123 
Polypodium, 80, 124-126; albidulum, 66; 
albobrunneum, 124; alternifolium, 120, 

121; arnorphum, 127; australe, 85; cam- 
bricum, 85; costatum, 121; discolor, 67; 
hesperium. 35, 126; heterophyllum, 102; 
kalbreyeri, 66; leptophyllum , 101-103; 
marginale, 102; sect. Marginaria, 66; 
mathewsii, 65, 66; montease, 127, 167, 168; 
nigrescens, 120, 121; sect. Polypodium, 66; 
plukenetii, 120, 121; princeps, 118; quad- 
ripartitum, 106; scolopendria, 120, 121; 
sechellarum, 124; trifurcatum, 127; varium, 
102; virginianum, 167; vuigare, 85, 142, var. 
hesperium, 35, var. serratum, 128 
Polystichum, 35, 116, 124; kruckebergii, 167; 

scopulinum, 167; aquarrosum, 142 
Proferea, 120 
Proneplirium, 120 
Psilotum, 32; nudum, 30-32, 161 
Pailotum nudum New to Arizona, 30 
Psilotum nudum Spreading Northward, 161 
Pteridium, 25; aquilinum, 13, 25, 26, 142 
The Pteridophyta of Taiwan-5. Hymeno- 

phyUaceae-.Trichomanes (rev.), 87 
Pteris ensiformis, 142; multifida, 142; quad- 
riaurita, 125; serrulata, 142; vittata, 128; 
wallichiana, 110 
Ptychophyllum magellanicum, 105 
Puri, H. S. Indian Pteridophytes Used in 

Folk Remedies, 137 
Pyrrosia, 118; costata, 119, 121; grandissima, 
119; lingua, 118; princeps, 118, 119; 
splendena, 119 
Pyrrosia princeps, a Fern New to Cultivation, 

Reed, Clyde F, Bibliography to Floras of 

Southeast Asia (rev.), 39 
A Reinvestigation of the Morphology of 

Hypodematium crenatum, 107 

Report of: Auditing Committee, 45; Fern 

Spore Exchange, 46; Judge of Elections, 45; 

President, 40; Secretary, 41; Treasurer, 42 

Reviews of Tropical African Pteridophyta, 

1 (rev.), 123 
Reviews: Bibliography to Floras of Southeast 
Asia, 39; Chromosomes and Speciation in 
Ferns, 39; A Commentary on Some Type 

Speciiiiens of Ferns in the Herbarium of 
K. B. Presl, 119; Fern Hybridizing at the 
University of Leeds, 86; Flora Palaestina, 
Part One. Text, 127; The Illustrated Flora 
of Illinois: Ferns, 32; Karyological Studies 
on Aspleniaceae 1. Karyotypes of Three 
Species in Asplenium, 86; A New Name for 
a Species of Polypodium from Northwestern 
North America, 126; Pacific Northwest 
Ferns and Their Allies, 167; The Pterido- 
phyta of Taiwan-5. Hymenophyllaceae: 
Trichomanes, 87; Reviews of Tropical 
African Pteridophyta-1, 123; A Revised 

Flora of Malaya, vol. II-Ferns of Malaya, 
ed. 2, 38; Vascular Plants of the Pacific 
Northwest, part 1, 34; The Victorian Fern 
Craze, 84; Welsh Ferns, Clubmosses, 
Quillworts, and Horsetails, A Descriptive 
Handbook, 84 

A Revised Flora of Malaya, vol. II— Ferns 
of Malaya, ed. 2 (rev.), 38 

The Rhizome Scales of Platycerium, 144 

Salvinia aiiriculata, 125; minima, 125; 
rotiindifolia, 124, 125 

Salviniaceae, 123 

Schelpe, E. A. C. L. E. Reviews of Tropical 
African Pteridophyta, 1 (rev.), 123 

Selagineila, 6, 35, 142, 169; martensii, 1-3, 5, 
6; willdenovii, 5 

Setzer, R. B. Kinds of Cystopteris in Cah- 
fornia, 164 

Stelar Anatomy of Six Species of Lycopodium, 

Stem Cross-section Prints Aid in Identifymg 

Horsetails, 161 
Stenochlaena cumingii, 123; laurifoha, 12-, 

123; palustris, 123 
A Survey of Some Morphological Features of 
the Genus Elaphoglossum in Costa Rica, 73 
Tatuno, S. Karyological Studies on Aspleni- 
aceae 1. Karyotypes of Three Species m 
Asplenium (rev.), 86. 
Taxonomic Notes on Ferns, IV, 103 
Taylor, T. M. C. Pacific Northwest Ferns 

and Their Allies (rev.), 167 
Tectaria, 34, 116; polymorpha, 142 
Terminology of the Sporangial Structures 

Equisetum, 162 < rvo 

Thelypteris, 24, 34. 124; dentata. 13, 14, 23; 
sect. Glaphyropteris, 103; megaphylla, 
120; nevadenais, 168; palustris, 24, 128 
Thompson, J. W. (see C. L. Hitchcock) 
Trichomanes, 87, 124; scandens, 87 
Two New State Records for Isoetes m Ken- 
tucky, 30 ^, . f 
Vascular Organization of the Rhizome of 

Cibotium barometz, 68 - 

Vascular Plants of the Pacific Northwest, 

Part 1 (rev.), 34 

The Victorian Fern Craze (rev.), 84 

Vittariaceae, 123 

Wade. A. E. (see H. A. Hyde) 

Webster, T. R. Aberrant Leaves on Angle- 
Shoots of Selagineila martensii Spring, 1 

Welsh Ferns, Clubmosses, Quillworts, and 
Horsetails, A Descriptive Handbook (rev.), 

Woodsia, 33, 34; ilvensis, 34; obtusa, 34 

Woodwardia chamissoi, 36; fimbriata, 
paradoxa, 36; radicans, 36 

Xiphopteris, 123, 124; albobrunnea. 
villosissima subsp. subpinnata, 124 

Zoharv, M. Flora Palaestina, Part 
Text (rev.), 127 




Page 105, line 3: For "B. araucana^^ read ^'B. araucanum .^ ^ 
Page 125, line 8: For ^'A. auriculata^' read ^^>S. aiiriculata.^^ 
Page 127, line 8: For ''1166'' read ''11667 J' 


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197 1 



VouTME 61, Number 1, Pages 1-48, Issued March 23, 1971 

A Study of Azolla pinnata R. Brown A. Bweet and L. V. Hills 1 

A New Variety of Lindsaea odorata Roxb- from Darjeeling in the 
Himalayas T. Sen and U. Sen 14 

The Proper Disposition of Meniscium macrophyllum Ktuize 

C. V. Morton 17 

The Thelypteris normalis Complex in the Southeastern United States 

A. Reid Smith 21 

Asplenium pinnatifidum 

ecord for Indiana 
Gerald J. Gastony 32 

A Dwarf Ostrich Fern H. Lou Gibson 35 

Shorter Notes: Niphidiiun longifolium, a Necessary New Combination; 

Bulbous Adder's-tongue Common in Louisiana 37 

Recent Fern Literature ^^ 

American Fern Society : Report on the 1970 Fern Foray 44 

Volume 61, Number 2, Pages 49-96, Issued June 24, 1971 

Lindsaea (Schizoloma) ensifolia Swartz in Hawaii. W. H. Wagner, Jr. 49 

The Fern Collections in Some European Herbaria, VII 

C. V. Morton 59 

An Appendageless Psilotum. Introduction to Aerial Shoot Morphology 

Albert S. Rouffa 75 

Vascularization of Fern Leaves 

Robert C. Lommasson and C. H. Young, Jr. 87 

Shorter Note: A Wrongly Localized Species of Pyrrosia 94 


Recent Fern Literature 

Volume 61, Number 3, Packs 97-144, Issued November 15, 1971 

The Fern Vegetation of Aldabra Atoll F. R. Fosbkkg 97 

Leaf Epidermal Studies in Marsilea 

John T. Mickel and Frank V. Votava IOI 

The American Species of Plagiogyria sect, Carinatae 

David B. Leujnoicu 110 

The Genus Stenochlaena J. Smith with Description of a New Species 

R. E. HOLTTUM 119 

The Variation in Spore Size and Germination in Dryopteris Taia 

Dean P. Whittier and W. H. Wagner, Jr. ] 23 

The Gametophytes of Natural Hybrids in the Fern Genus Pellaea 

Thomas R. Pray 128 


Schemochromic Blue Leaf-surfaces of Selaginella 

Denis L. Fox and James R. Wells 137 

Shorter Notes : Asplemum kamtchatkanum Gilbert, a Dubious Eastern 
As'atic Fern ; A New Locality for Psilotum nudum in Sonora, Mexico . . 140 

Recent Fern Lterature 142 

Voi,T-ME 61. NrMMER 4, Pages 145-192, Issfed Dkcember 17, 1971 


The Genus Grammatosorus C. V. Morton 145 

The Genus Selaginella in Baja California, Mexico Ira L. Wiggins 149 

Notes on the Ferns of Dominica and St. Vincent, II 

C. V. Morton and David B. Lellingeu 161 

Two New Tree Ferns from Costa Rica Luis Diego G<5mez 16t) 

Comparative Habitat Requirements for Spore Germination and Pro- 

thallial Growth of Three Ferns in Southeastern Michigan 

RoYCE H. Hill 171 

The Gametophyte of Thelypteris erubescens Lenktte R. Atkinson 183 

Shorter Notes; A Recent Find of Isoetes in Louisiana; Jamesonia 

alstonii in Oaxaca, Mexico '86 

Recent Fern Literature 188 

Index to Volume 61 ^88 





Vol. 71 C6/ a Januart-Maech, 1971 No. 1 

American Jfern journal 










A Study of AxoUa piimatft R, Brown. . ^. ^ ^.A. Sweet and L. V. Hiixs 1 

A New Variety of lindsaea odorata Roxb. from Darjeeling in ^ 
Bhnalajras T. Sen and U. Sen 14 

Ibe Proper Disposition of Menisctom macrophylhun Konze 

a V. MOBTON 17 

Tlie *^ elypteris aonnalis Complex in the Southeastern United States 

A. Reid Smith 21 

Aspleniom pinnatifkltim X trichomanes — A New Record for Indiana 

Gekald J. Gastont 32 

A Dwarf Ostrich Fern H- Lot; Gibson 35 

Shorter Notes: Niphidium longifolinm, a Necessary New Combination; 
BiUbous Adder*s- tongue Common in Louisiana. 37 

Recwit Fern Literature *J 

American Fern Society : Report on the 1970 Fern Foray ,.,. ti 

Ai , •::"^| 

(Efie American jftvn ^otietp 

+ * -f^pi 

Council for X971 

Warren H, Waqneh, Jr., Department of Botany, Univeraity of Michigan, 
Ann Arbor, Michigan 48104. President 

JfOHN T. MiCKEL, New York Botanical Garden, Bronx Park, Bronx, New 
York. 10458. Vice^Prestdent 

Richard L, Hauke, Department of Botany, University of Rhode Island, 

Kingston, Rhode Island 02881 Secretary 

LeRot K. Henry, Diviaion of Plants, Carnegie Museum, Pittsburgh, Penn- 
sylvania 15213. Treasurer 

David B. Lellinger, Smithsonian' Institution, Washington, D. C. 20560. 

Ediior -in-Chief 


r ■ 

Rational ^ocietp B^epte^tentatibei^ 

Warren IL Wagner, Jr., University of Michigan A.A.A.S. Representative 
RoLLA M. Teton, Jr., Harvard University A.LB.S. Representative 

Simttitan ftxn Journal 


David B, Lellinger Smithsonian Institution, Washington, D. C. 20560, 

C. V. Morton Smithsonian Institution, Washington, D. C. 20560. 

RoLLA M. Tryon, Jr. 

Gray Herbarium Harvard University, Cambridge, Mass. 02138. 
Ira L. Wiggins . . . .Dudley Herbarium, Stanford University, Stanford, Calif. 

An illustrated quarterly devoted to the general study of ferns, owned by 
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Matter for publication should be addressed to the Editor-in-Chief. 

Changes of address, applications for membership, subscriptions, order? 
for back numbers should be addressed to the Treasurer. 

Subscriptions $5.50 gross, $5.00 net (agency fee $0.50); sent free to mem- 
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ftporc Cxftanae 

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' Gifts and bequests to the Society enable it to expand its services to mem- 
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American Jfern fottrnal 

VoL.-Tl C ^ i) ^ January-March, 1971 No. 1 

A Study of AzoUa pinnata R. Brown 


A. Sweet and L. V. Hills^ 

A studj^ was undertaken to ascertain the morphologic variability 
of the vegetative and fruiting structures of plants within the 
Azolla pinnata complex. Plants of this group have been commonly 
referred to three species, A. pinnata j A. africana^ and A, imhricata. 
Our observations indicate that only one species is present, although 
two varieties based on vegetative morphology can be established. 
The synonymy given by Christensen (1905, p. 148) forms the 
basis of our synonymy for A. piniiata. In addition to Christensen, 
the original diagnoses and the work of Baker (1887) and Shen 
(1961) were consulted in working out the taxonomy of this group. 
Anatomical studies have been done on these plants by Griffth 
(1845), Strasburger (1873), Sud (1934), Rao (193G), Dehnalsy 
(1958), and Bonnet (1957). :\loore (1969) gives a general review 
of the literature on Azolla and indicates its use in agriculture. 

A total of 85 collections were examined, 70 in detail When 
available, at least 10 plants per collection were examined under a 
binocular microscope. Wetting was sometimes used to bring out 
detail. Alassulae were placed in water under a vacuum for two 
hours to remove air bubbles. Following this, they were mounted in 
polyvinyl alcohol (Elvanol solution) on a cover slip which, when 
dry, was affixed to a slide with Canada balsam. The megaspore 
complexes were treated with acetolysis solution for 5-10 minutes 
at 200''F, washed, and then oxidized with Shultze solution (HXOs 

^The authors would like to acknowledge the co-operation of herbaria that 
contributed to this study by lending their oollectiorus. Scanning electron photo? 
were taken by .Air. J. Bloor. Mr. L. Johnson helped with the photography and 
Mr. P. Wolterri with the drafting. The manuscript w^.^ typed by Miss N. 
Fraser. Expenses were covered by a National Research Council Grant-in- 
Aid of Research to L. V. H. 

Volume 70, No. 4, of the Journal, pp. 12^172, was issued Jan. 6, 1971. 

2 American Fern Journal 

-j- KCIO3). ATounting followed the same procedure as with the 


The definition of new terms and terms which have been used as 
applied to Azolla in conflicting ways are as follows: 

Trichomes. — The spine-like projections arising from the angles 
between the contact areas on the proximal siu'face of the massulae, 
having vacuoles continuous with those of the massulae (Shen, 
1961). The restriction of trichomes to the proximal surface and the 
distribution of vacuoles differentiates trichomes from glochidia 
which occur in the species of section Azolla, 

Megaspore complex. — The composite structure from which the 
female gametophyte develops. 

Cap, — The remnant of the indusium persisting as a total or 
partial covering over the floats (Figs. 1^2). 

Supraspore (new term). — The apical region of the megaspore 

complex (Fig. 1) consisting of: The columella or apical extension 

of the perine forming a central conical projection (Fig. 6); the 

columnar filaments or fibrils arising from the apex of the columella 

and spreading over the surface of thefupper floats; and the floats 
{Figs, 1,2). 

Collar, — The region intermediate between the supraspore and 
the infraspore formed by a thickening of the perine. 

Infraspore (new term). — The distal area of the megaspore com- 
plex (Fig. 1) consisting of the exine or wall of the megaspore or 


germinal body (Figs. 3, 5, 20) ; and the perine or wall layer formed 
exterior to and free from the exine {Figs. 1,2). 

Inperine (new term). — The inner, homogeneous or vacuolate 
zone of the perine located adjacent to the exine {Figs. 5, 20). 

Experine (new term). — The outer, sculptured zone of the perine. 

Experine 1 (new term) .—The inner zone of the experine with a 
vacuolate or pillar-like character which acts as a foot layer for the 
outer elements of sculpturing {Figs. 5, 20). 

Experine 2 (new term). — The outer zone of the experine com- 
posed of the larger elements of sculpturing {Figs. 5, 20). 


The subdivision given for the infraspore agrees in principle with 
that of Strasburger (1873), Rao (1936), Dehnalsy (1958), Bonnet 
(1957), Kempf (1969) and Snead (1969), but differs from that of 
Hall (1968), who considers only the outer fibrous layer found on 
some fossil forms of AzoUa to be perisporial material. He includes 
our experine and inperine as part of the megaspore exine. 
AzoLLA PiNNATA R. Brown, Prodr. Fl. Nov, Holl. 167. 1810. 

Plants small, floating aquatics with bilobed leaves. Pubescence 
prominent on upper surface of dorsal leaf lobe and on stem near 
leaf base, sometimes extending over entire internodal area, and 
less commonly onto margin of ventral leaf lobe. Ventral leaf lobes 
(Fig. 15) composed of single layer of cells, 1.5-2 times larger than 
dorsal leaf lobes. Two or more leaves before first lateral branch, 
then one leaf between succeeding branches {Fig. 25). 

^Megaspore complex 425-660 /x (avg. 560 ju) long. Caps 200- 
250 /x long, 3^-% projecting above supraspore {Figs. 1^2), Gen- 
erally 9 floats symmetrically arranged in groups of 3; each group 
containing 1 diamond-shaped upper float 1.5-2 times the size of 
the 2 trapezoidal lower floats {Fig. 21). Sometimes one extra small 
lower float {Figs. 1,2). Upper floats covered by columnar filaments. 
Inperine and experine apparent in columella {Fig, 6). Expansion of 
mperine at top of infraspore forming indistinct coflar {Fig. 6). 

Infraspore 225-450 fi (avg. 310 ju) wide, composing l^i^H of 
megaspore complex. Trilete rays }4 length of megaspore radius; 
exine perforate, 7 n thick {Fig. 3). Ferine 16 m thick with homo- 
geneous inperine 6 /z thick {Figs. 5, 20) ; experine 1 of irregular 
pillar-like structures, 1-2 m thick {Fig. 20); and experine 2 of 
irregular branching and fusing baculae 2-5/1 in diameter, 8-9 jit 
thick {Figs. 5, 20). Projecting from experine, convoluted pro- 
longations 12-14 /x long, consisting of a hollow central cylinder 
10-30 11 in diameter arising from inperine to which are fused 
swollen baculae {Figs. 4j ^0). Prolongation density variable 
{Ftgs. 7-9), Surface filaments rai^e or lacking, 

Aficrosporangia containing 3-8 massulae each having a convex 
distal and a pyramidal proximal surface {Fig. 13). INIassuIae 135- 
210 M (avg. 190 m) long, 150 y. wide, and 100 m deep. Trichomes 
2-10, usually 4-5 in number, 45-150 ii long, 4-20 ju wide at base; 
varying in shape from threadlike to stocky {Figs. 11, 13). Spines, 
structurally resembling short trichomes, up to 20 m long at the 
corners of some massulae {Fig, 11) or less frequently around the 
entire equatorial margin {Fig. 12). Massulae surface granular or 
finely vacuolate. Microspores 18 /i in diameter. 

American Fern Journal 

Volume 71, Plate 1 


s por e 



Details of Megaspore Complexes 




134, 135 (Presented by direction of J. J. Bennett, 1876) (E!). 
Range: As in Plate 4, see also p. 7). 

The size of the )nogas])ore romplex and component structures 
was found to be relatively constant within individual collections, 
but the total inter-collection variability was large. In particular, 
megaspores complexes from Africa were notably larger (avg. 
600^) than those from other regions (avg. 475 ju). As this size 
difference was not correlated with any distinctive structural fea- 
tures either within the megaspore complex or other plant parts, 
we believe it best not to use it to establish a distinct taxonomic 



group. Inter-collection variation in the density of the prolongations 
occurred randomly with respect to geographic distribution; there- 
fore, it is not taxonomically useful. 

In 5-10% of the specimens one additional float appears either 
by division of one of the lower floats (Figs. 2, 22)^ or as an extra 
small float at the base of the supraspore either between two 

floats {Fig. 23) j or below and partially replacing one float (Figs, 
If 24), The occurrence of floats supplementary to the basic number 
of nine may be used in attempts to work out the phylogeny of 
Azolla (see Hills and Gopal, 1967). One double megaspore complex 
was observed {Fig. 10) w^here two infraspores were fused together, 
the supraspores being absent (see also Strasburger, 1893, Fig, 100). 
The greatest development of spines on massulae was seen on 
specimens from Africa. Specimens from AustraUa and New Guinea 
had spines only at the corners of lenticular-shaped massulae. Al- 
though not all massulae from Africa, Austraha and New Guinea 
had spines, this was the only morphologic variation which cor- 
related in part wdth specific plant types. In the future this feature 

Fig. 1. Scanning electron photos of megaspore complexes with supple-. 



infraspores illustrating varlltion in prolongation density. fig. 10. 

Fused infraspores. 

Ameuican Fern Journal 

Volume 71, Plate 2 

^p'A'-^-^-"^ '- '^.' V^^-"^^*^-"-^ "^ ^.T^ 

T-l^ 1. 

^m^.-^ L^^VHW^^r^J >-T~Fr^x T^V ■ I 

- »■ ■ V.Vf 




■ ■ --^^^^ 







■ ^ 











Details of Massulae and Leaves 


should be noted, as it may be diagnostic. 

Branched trichomes (Fig. 14) were observed by Rao (1936). The 
abundance of branching is variable, occurring in about 10% of the 
trichomes. Within the same collection or even in massulae from the 
same microsporangia some massulae have branched trichomes, 
whereas others lack them; hence the presence or absence of 
branched trichomes is not a reliable taxonomic character. 

Morphologic variability in the fruiting structures cannot be 
used to subdivide A. pmnata into varieties. However, on the basis 


AzoLLA PiNNATA R. Brown var, pinnata 
Azolla africana Desv. Ann. Soc. Lhm. Paris 6: 270. 1827. 
Azolla guineensis Schum. Beskrivelse af Guineiske planter. 462. 
1827 [a preprint]. See also Dansk Vid. Selsk. Skr. Natur. Mat. 


138. 1887. 


Plants 1.5-2.5 cm long, 1-1.5 cm wide, with dominant main 
axis and pinnate appearance (Fig. 18). Dorsal leaf lobes 0.8-1.9 
mm long, 0.5-1.10 mm w^ide, length-width ratio 1.4-2.1, imbricate 
to highly imbricate (may be non-imbricate), relatively thick, 
curled both longitudinally and transversely, elongate, acute at the 
apex, with a hyaline margin of irregular width formed by up to 
four rows of elongate cells, the tips protrudhig to give a serrated 
margin (Fig. 19), Echinate pubescence on basal portion of dorsal 
leaf lobe grading to sometimes dense, irregularly arranged, conical 
pubescence on upper part, this sometimes accompanied by longi- 
tudinal striations. Leaves 4-16 (avg. 8) before the first branch. 

Range: Austraha, New Guinea, Africa and Madagascar. Col- 
lection 30 {PI 4) from Thailand is most similar to plants from 
New Guinea and may represent a population introduced for agri- 
cultural purposes; otherwise it is disjunct in its occurrence). 

COLLECTION'S SEEN: See page 11 and Plate 4- 
Azolla pinnata var. imbricata (Roxb.) Bonap. Notes Pterid. 

7: 130. 1918. 

Fig. U. Equatorial view of massulae showing end spines, trichomes 


SPINES. Fig. 13. S.E.M. photo of oblique proximal view of massulae. 
Fig. 14. Trichome with bifurcated tip. Fig. 15. Ventral leaf lobe. Fig. 
16. Plant of var, imbricata. Fig. 17. Dorsal leaf lobes of var. imbricata. 
Fig. 18. Pl.\nt of var. pinnata. Fig. 19. Dorsal leaf lobes of var. pinnata. 

American Fern Journal 

Volume 71, Plate 3 



experine 2 

experine 1 









p 10 






1 3 5 7 9 n 13 15 






Morphology and Distribution of A. pinnata. 

Fig. 20. Schematic diagram of megaspore complex wall morphology. 
Fig, 21. Relative size and shape of floats. Fig. 22. Left lower float 
DivujED. Fig. 23. Additional float between pair of lower floats. Fig. 
24. Additional float below one lower float. Fig. 25. Schematic repre- 
sentation- of leaf-branch pattern. Fig. 26. Distribution of the number 
OF leaves before the first lateral branch. Fig. 27. Distribution of 




Salvima imbricata Roxb. ex. Griff. Calcutta J. Nat, Hist. 4: 
469. 1844. 

Azolla imbricata (Roxb.) Nakai, Bot. Mag. Tokyo 39: 185. 1925. 

Plants 1-1.5 cm diameter. Main stem not dominant, but with 
dichotomous-like type of branching {Fig. 16), Dorsal leaf lobes 
0.8-1.5 mm long, 0.6-1.0 mm wide, length-width ratio 1.1-1.5, 
generally non-imbricate to slightly imbricate, thin, flat, ovate 
to trapezoidal, rounded to subrounded at the apex, with a hyaline 
margin of regular w^idth formed by 2-5 rows of nearly equidi- 
mensional cells {Fig, 17), Echinate pubescence on basal portion of 
dorsal leaf lobe grading to evenly distributed conical pubescence 
on the upper part {Fig. 17). Leaves 2-8 (avg. 4-5) before the 
first lateral branch. 


Range: India, Nepal, Ceylon, Burma, Southeast Asia, Philip- 
pines, China, and Japan; one disjunct collection from Africa. 
Plants intermediate between var. pinnata and var. imbricata are 
found in Australia, New Guinea, Africa and India. 

Collections seen: See page 11 and Plate 4- 

Azolla pinnata var. imbricata is distinguished by the plants 
being generally smaller and more circular in shape, with rounded 
to subrounded, ovate to trapezoidal dorsal leaf lobes, these with 
a regular hyaline margin of nearly equidimensional cells and 
regularly distributed conical pubescence. 

The trivial name imbricata was an unfortunate choice by 
Roxburgh (1844), who compared them with the non-imbricate 
Salvima. To him the Azolla would seem imbricate, as the ventral 
leaf lobes are overlapping and the dorsal leaf lobes are never ^-idely 
Spaced. However, we have found these plants to be non-imbricate 
compared to those of var. pinnata. 

The distinctiveness of the leaf-branch pattern in different species 
of Azolla was first suggested by Strasburger (1S73). Bonnet (1957) 
states that in A. pinnata 3-9 leaves precede the first branch and 
the number of leaves before the first lateral branch increases 
toward the apex of the plant. We observed 2-1 G leaves and no 
consistent increase or decrease towards the apex. Fiijure B6 shows 
the distribution of average leaf numbers before the first branch, 
based on at least 10 counts per collection. Although in general 

American Fekn Journal 

Volume 71, Plate 4 

Characters and Geogr 



fewer leaves occur before the first branch in var. tmhricala than in 
var. ptnualUy leaf-branch pattern is a more useful criterion for 
separating species of Azolla (Bonnet, 1957) fhan for separating 
varieties within A. pinnata. 

Figure 27 shows the distribution of length-width ratio for the 
dorsal leaf lobes, based on at least five mature specimens per 
collection. This ratio, if considered in conjunction with other 
criteria, is useful in distinguishing the two varieties. 

_ * 

The leaf-branch pattern, the apical shape, the degree of imbrica- 
tion, the type of pubescence, and the length-width ratio of the 
dorsal leaf lobe are symbolically represented with respect to geo- 
graphical distribution in Flaie 4- In the preceduig descriptions 
and in Plate 4 the term *'non-imbricate'' describes leaves in which 



there is no overlap; '^highly imbricate" when they overlap for at 
least 2/3 of their length, and ''imbricate" when the overlap is 
intermediate. ''Regular conical" indicates evenly distributed pu- 
bescence whose length is less than twice the width and ''elongate 
conical" indicates irregularly distributed conical pubescence ac- 
companied by longitudinal striations. 




2673 (US); 12, L. Brass 5843 (C); 13, A. Harre 270 (F); 14, J. Brown-Lester in 



(UC); 21, A. Peter 44388 (UC); 23, R. Schlechter 12033 (M); 24, H. Soyaux, 
24/11/1874 (M); 25, H. M. Richards 8313; 26, U. B. Drummond & A. J. 
Cookson 6272; 27, /. M. Hildebrandt 3065 (M) ; 28, G. F. S. Elliot 1798 (E) ; 30, 
Th. Sorensm, K. Larsen & B. Hansen 547 (C). Examined but not plotted: 
N. H. W. Brown (E), Gower Collection, River Quenso (E). 

Var. imbbicata: 17, P. Cadjoe 983 (M); 32, Em. Bodinier 1427 (E); 33, 
M. S. Clemens, 3/1907 (M); 34, A. Loher II49 (M, US); 35, H. M. Curran 
11624 (C); 36, A. M. Evans 54O (TENN); 37, G. Murata 27129 (US); 38, 
SieboW in 1842 (M); 39, T. Nakano in 6/1896; 40, M. Hutch 17883 (E); 
41, L. H. Bailey in 7/27/1917 (US); 42, A. Henry 3977 (CAL); 43, H. C. Cheo 
501 (E); 44, Kweichow Expedition 4220 (C); 45, H. Smith 1518 (C); 46, Th. 
Sorensen, K. Larsen & B. Hansen 1829 (C); 47, Urupai on 2/21/1969; 48, 
/. Sinclair on 12/12/1948 (E); 49, J. Sinclair on 14/11/1950 (E); 50, Fer- 
ffuson (M); 51, Thwailes C. P. [Ceylon Plants] I42I (E); 52, Marshall Ward 

12 American Ferx Journal 

in 10/1880 (E); 53, A. 0. Brodie in 9/186? (E); 55, C. E. C. Fischer Jfi27 
(CAL) ; 56, Dr. Wight in 1831 (E) ; 57, Shaik Mo Kim in 1896 (CAL) ; 58, P. 
M. Debbarman 6U (CAL) ; 59, S. Kurz 3194 (CAL) ; 60, J. D. H. (C) ; 61, J. J. 
Wood 187 (CAL); 62, C.Maries 1890 (CAL); 63, H.H. Johnston on 1/27/1896 
(E) ; 64, 0. Polunin, W. R. Sykes & L. H. J. Williams, 1822 (E) ; 65, Stainton, 
Sykes & Williams 6952 (E); 66, D. H. Nicolson 2911 (US); 67, Stainton, 
Sykes & Williams 2546 (CAL, E) ; 68, East India, Dr. Wight (E) ; 69, H. 
Collett, 11/1890 (CAL); 70, G. Watt 5090 (CAL). Examined but not plotted: 
M. Vanoverbergh 3226 (BKL) ; R. R. Stewart 1359 (BKL) ; Gower Collection (E) ; 
Wight 5 (C). 

Intermediate types: 3, J. Camfield in 4/1897 (US); 6, N. Michael 1431 (E); 



(M); 31, G. H. Vachell on 17/7/1830 (E); 54, Stocks, Law (C). 

The distribution of the two varieties presents an interesting 
problem. The close similarity of the disjunct populations of var. 
pinnata indicates the populations were once connected. INIega- 
spore complexes similar to those of A . pinnata in Upper Pliocene- 
Lower Pleistocene strata of Russia (Nikitin, 1957; Dorofeve, 
1963) indicate the species was once much more widespread. 
It is therefore reasonable to assume that the separation of popu-. 
lations of var. pinnata may have occurred as the result of glaciation 
during the Pleistocene (ca. last 3 million years) . 

Plants typical of var. pinnata were not seen in collections from 
India and Ceylon. However, some plants from these regions had 
notably higher leaf numbers before the first branch than usual is 
for var. imbricata. Other plants had pubescence more like that of 
var. pinnata. This may suggest a former relic population of var. 
pinnata in India and/or Ceylon which has influenced the character 
of, and has been absorbed into; the present population. 

Attempts were made to relate the distribution of the two vari- 
eties to generalized plots of variations in temperature and light 
intensity. No direct relationship between these factors and the 
distribution pattern was found. It was noted that var. imbricata 
occurs m more densely populated areas, suggesting that some 
agricultural practice may influence the distribution. However, the 
succession of plants from Australia to Southeast Asia appears to 
reflect a clinal development, and therefore agriculture is probably 
not the controlling factor. 


Future paleobotanic studies will unquestionably add to our 
knowledge of the former distribution of the species. Tlie answer lo 
the relationship between the two varieties and the reasons for the 
distinct geographic ranges will best be answered by the study of 
the varieties themselves. Future collections should be more specific 
as to the character of the collecting site and its ecological setting. 

Literature Cited 

* ■ 

Baker, J. G. 1S87. Handbook of the fern allies. George Bell, London. 
Bonnet, A. L. M. 1957. Contribution k V6iude des Hydopt^rid^es. III. 

Recherches sur AzoUa filiculoides Laink. Rev. Cytol, Biol. A'^get. 

18: 1-88. 

Christensen, C. 1905-6. Index Filicum. Hagerup, Copenhagen. 

Delmalsy, p. 1958. Nouvelles recherches siir le sporophyte d^AzoIla. La 

Cellule 59 : 235-268. 
Dorofeev, p. I. 1963. The Tertiary floras of Western Siberia. Akad. Nauk 

SSSR, Bot. Inst. Komarov, Leningrad. 
Griffith, W. 1845. On Azolla and Salvinia. Calcutta J. Nat. Hi^t. 5: 227-273, 
Hall, J. W. and N. P. Swanson 1968. Studies of fossil AzoIIa: Azolla mon- 

tana, a Cretaceous niegaspore with many small floats. Amer. J. 

Bot. 55:1055-1061. 
Hills, L. V. and B. Gopal. 1967. Azolla primaeva and its phylogenetic 

significance. Canad. J. Bot. 45: 1179-1191, 
Kempf, E. K. 1969. Elektronenmikroskopie der Sporodermis von kanozoischen 

Megasporen der Wasserfarn-Gattung Azolla. Palaont. Zeitschr. 43: 

Moore, A, W. 1969. Azolla: Biology and agronomic significance. Bot. Rev. 

NiKiTiN, P. A. 1957. Pliocene and Quaternary floras from Voronezh District. 

Acad. Nauk SSSR, Bot. Inst., Moscow-Leningrad. 
Rao, H. S. 1936. The structure and life history of Azolla pinnata. R, Br. with 

remarks on the fossil history of the Hydropterideae. Proc. Indian 

Acad. Sci. 2: 175-200. 
Shen, E. Y. F. 1961. Concerning Azolla imbricata. Amer. Fern J. 51 : 151-155. 
Snead, R. G. 1969. Microflora! diagnosis of the Cretaceous-Tertiary bound- 
ary, Central Alberta. Res. Coun. Alberta Bull. 25. 
Strasburger, E. 1873. Ueber Azolla. Friedrich von Zezschwitz, Jena. 
SuD, S. R. 1934. A preliminary note on the study of Azolla pinnata R. Br. 

J. Indian Bot. Soc. 13: 189-197. 

Department of Geology, University of Calgary, Calgary 
44, Alberta, Canada. 

14 • Americax Fern Journal 

A New Variety of Lindsaea odorata Roxb 
from Darjeeling in the Himalayas 

T. Sen and U. Sen 


During our field studies in Darjeeling in 1965 and 1967, we 
discovered some Lindsaea specimens that were strikingly different 
from the species of Lindsaea familiar to us. The plants resemble 
Lindsaea odorata Roxb.," which is also found in Darjeeling. How- 
ever, close examination showed that they are quite distinct: they 
are smaller and have short branched foliar hairs, but lack multi- 
cellular long hairs and internal endodermis in the rhizome. These 
plants thus represent an undescribed taxon. 

Lindsaea odorata var. darjeelingensis Sen & Sen, var. nov. 

A var. odorala differt: frondibus multo minoribus, usque ad 
lo cm longis et 1.7 cm latis; paleis rhizomatis latitudine basi ex 
celluhs 4-7 constantibus; frondibus juvenilibus pilis brevibus 
ramosis praeditis; endodermide interiore stelis rhizomatis carente. 

Rhizome creeping, short, slender, ca 2 mm in diameter; scales 
deep brown, acuminate, up to 7 cells broad at the base; vascular 
cylinder an amphiphloic solenostele but lacking an internal 
endodermis. Fronds simply pinnate, to ca 15 cm long, 1.7 cm 
wide; petiole slender, shallowly grooved on the adaxial surface, 

round below, to 4 cm long. 

short-stalked (0.5 mm 


veins immersed 

the baslscopic margin bearing 3 or 4 branches, each 1- or 2-forked, 

1-4 ^ ^^^ 

fronds with smaU, 2- or 3-celi?d 

3-celled branched glandular hairs; sporangia globose; annulus of 
io~Zo cells; stomium 8-12 cells; paraphyses present with sporangia; 
spores bilateral, 52 m X 38 /x X 33 n, enclosed by a thin perine, 
this cracking off on acetolysis. 

Type: Darjeeling, on way to Birch Hill, Sen & Sen 505 (holo- 


Profp.tJ'f P «^f Professor R. E. Holttnm for the Latin diagnosis, ajid 
iToiessor b. P. Sen for his encouragement 

v.hMh^Z?^''T\ -^- ^^^' ^?^'^^ ^"^^ «^"'«^" this to be the correct name for 
What ha^ alttay.s,but incorrectly, been called Lindsaea cuUrata (Willd.) Swartz. 

American Fern Journal 

Volume 71, Plate 5 


Fig. 2. Rhizome scale, X 53. Fig. 3. Pinna, X 5. Fig. 4. Unbranched 


view of epidermal cells on the abaxial side of lamtna, x 87. fig. 7. 

Transverse section of the rhtzome showing the absence of inner 
ENDODERMis, X 300. FiG. 8. Spore, X 435. 

16 American Fern Journal 

• Other specimens seen: Darjecling-Sudrapong Road, Sen & 
Sen 506] Raiigaroon Forest, Sen & Sen 507; Lebong, Sen & Sen 
.505; Darjeeling, (?. King BSI SSP, 5SU (CAL). 

This new variety somewhat resembles var. pallets Hook. (Sp. 
Fil 1: 204. i846) from Nepal ajid Sylhet in frond size, and yet 
differs in other characters. The features which distinguish and 
characterize var. pollens are: (i) the scales are less than 4 cells 
wide at the base, (ii) short branched hairs and multicellular hairs 
are seemingly absent even on the youngest fronds, (iii) the vas- 
cular cylinder of the rhizome has both the inner and outer endo- 
dermis, and (iv) the spores measure Gl ;x X 44 /i X 48 /i. We con- 
clude that var. darjeelingensis is a very distinct local variety of 
this wide ranging species. 

The differences among the three varieties of Lindsaea odorata 


may be summed up in the following key: 


Scales 2-3 or very rarely 4 cells broad at the base; short branched hairs ab- 
sent on the laminae; stele of the rhizome with both inner and outer 

Long multicellular hairs always present on young fronds; fronds ca 60 cm 

long and caj35mm wide at the broadest region; laminae thick; 

spores 64 M X 44 M X 43 /x. var. odorata 

Long multicellular hairs never occur even on the young fronds; fronds ca 

20 cm long and 24 mm wide at the broadest region; laminae 

thin; spores 61 ^ X 44 m X 48 ju. var. pallens 

Scales 4-7 cells broad at the base; short branched hairs always present on 

young laminae; stele of the rhizome with only the outer endodermis; 
long multicellular hairs never develop on the fronds; fronds ca 15 
cm long and 17 mm wide at the broadest region; spores 52 m X 38 m 
^ ^^ ^^' var. darjeelingensis 

Department of Botany, K^lyani University, Kalyani, 
West Bengal, India. 

Mentscium macrophyllum 17 

The Proper Disposition of Meniscium macrophyllum Kunze 

C. V. Morton 

In our paper ''The American Species of DryopteriSj Subgenus 
Meniscium'' (Bull. Torrey Bot. Club 65: 347-37G, 1938), Dr. 
Maxon and I concluded that Meniscium macrophyllum Kunze 
could not properly be referred to Dryopteris subg. Meniscium 
because some of the sporangia are borne on the leaf surfaces be- 
tween the veins, thus being ''acrostichoid." Consequently, we 
proposed to refer the species to Bolhitis, where the acrostichoid 
condition is the rule. However, I have never been really satisfied 
with this disposition, because this species resembles species of 
the subg. Meniscium much more than any Bolbitis species in most 



]My reexamination of the available specimens indicates that 
in some specimens the sporangia are borne only on the arcuate 
cross- veinlets and also, exceptionally in Meniscium, on the short, 
medial, outwardly excurrent veinlets also. Indeed it appears that in 
semifertile blades the sporangia are mostly on the excurrent 
veinlets (cf. Spruce S030). In some specimens the sporangia do 
extend out also onto the leaf surfaces, which is certainly a unique 
character in Meniscium but not sufficient to necessitate the 
reference of this species to a different genus, especially when the 
character is not shown by all specimens. Therefore, I now refer 
M, macrophyllum to Thelypteris subg. Meniscium: 


Menisciilitr n? 


IHeteroneiHron meniscioides Fee, Mem. Foug. 2: 93, /. 55. 1845. Type: Bahia, 

Brazil, Blanchet (not seen). I have seen two specimens collected 
by Blanchet in Bahia, numbers 222S and 2477, both at Kew, and 
these are very likely authentic and parts of the type collections. 

IPoecilopteris menhcioides (Fde) K. B. Presl, Epim. Bot. 175. 1849 [1851]. 

Phegopteris macrophylla (Kunze) Mett. Fil. Lech]. 2: 24. 1859, at least as 

to basionym. 
Nephrodhtm macrophyllum (Kunze) Keys. Pol. Cyath. Herb. Bung. 48. 1873. 
Dryopteris macrophylla (Kunze) C. Chr. Ind. Fil. Suppl. 1;35. 1913. 

IS American Fern Journal 

BoJbilis macrophylla (Kunze)' Maxon & Morton, Bull. Torrey Bot, Club 65: 

375. 193S. 

Additional specimens examined: 

Brazil: San Isidro, Goyaz, Pohl (BR). Limocero, Goyaz, Pohl (BR). Serra 
do Mar, Bahia, Luschnath' (BR). Ilheos, Bahia, Luschnath^ (BR). Without 
loraIitj% Riedel (BR, sterile). 

Suriname: Jodensavanne-AIapanne Creek, Suriname River, Lindcman 
4945 (U); Hekking 1212 (U). Blanche ]\Iarievallen, Nickerie River, Maas tt 
Tawjoeran 10987 (U). * • 

British Guiana: Appun 7S2 (K). 

In our discussion of this species, INIaxon and I decided that 
specimens with setose sporangia ought to be referred also to B. 


viacroiylla^ but I have now changed my mind about that also. 
The type of M. macrophylhim and other Brazilian specimens 
that I have recently examined (but not all of those cited in 1938 
have been reexamined) have the sporangia non-setovse. Those 
from Trinidad and Tobago and some of those from the Guianas 
have them conspicuously setose, and in a peculiar way. The setae, 
numbering from one to four, are exceedingly conspicuous and are 
borne on the annulus. In the other species of Menisdum with 
setose sporangia we stated that the setae are borne on the stalk 
of the sporangium. I am now unable to verify this in all cases. 
Sometimes the seta is on the sporangium stalk, but at other times 
it seems to be only from the receptacle rather than the sporangia, 


thus being technically a paraphysis. It may be that thCvSe setae 
are too easily detached and are separated from their sporangium 
readily in making a microscope slide. However this may be, it 
does seem that the character is still a usable one in separating 
those species with setose sporangia or paraphysate sori from those 
with glabrous sporangia and sori. Since I have used this char- 


acter of setose sporangia as a fundamental character in this group, 
it is not reasonable to allow both types within T. macrophylla^ 
especially since there is some geographic disjunction and some 
other characters of more debatable importance. It appears that 
in typical T, macrophylla the excurrent veinlet is well developed 

1 These collections may very well be isotypes, for the specimens distributed 
by Martins in his numbered sets were often collected by others, among them 
Luschnath. They are in any case topotypes and quite identical to Manias 363, 

IMeniscium macrophyllum 19 

in the fertile pinnae, and it bears sporangia. In the plants with 
setose sporangia the arcuate cross veinlets are nearly straight and 
the excurrent veinlet is generally reduced to a mere knob hardly 
long enough to bear sporangia. This character ought to be checked 
in the field. I do not have enough good fertile material. This plant 
with setose sporangia may be known as: 
Thelypteris guyanensis (Fee) Morton, comb. nov. 

Meniscium gvyanense Fee, Gen. Fil. 224. l^.'>2. Type: **IIabitat in sylvis 

paludosis, ad amnes Conana et Gabaret/' French Guiana, Leprieur 
in 1835 (originally determined as M. sorbifoUurn) (P?, not seen). 
The description, especially the statement that the sporangia are 
pilose, indicates that the name refers to this species, 

Acrostichum fendleri Baker, Journ. Bot. Brit. & For. 25: 100. 1887. Type: 

Trinidad, Fendler 88 (presumably K, not seen; isotypes NY, US). 

Meniscium oUgophyllum Hort. ex Baker, Ann. Bot. 5: 487. 1891. Lectotype: 

Hort. Linden, ex Herb. Moore (K). A specimen '*e caldario Linden*' 
in Brussels is probably an isolectotype (Weatherby photograph US). 
A later collection, doubtless from authentic material, is cult. Jan. 7, 
1871, ex Herb. R, Sims (K). Probably this name ought to be con- 
sidered as invalid, since it was not definitely accepted as a valid 
species by the publishing author, Baker. 

Leptochilus fendleri (Baker) C. Chr. Ind. Fil. 385. 1906. Based on Acrostichum 

fendleri Baker (not Meniscitim fendleri Eaton^). 

Dri/opleris anceps Maxon, Contr. U.S. Nat. Herb. 24: 62. 1922. Based on 

Kuntze (1891). 


I have not been able to reexamine all the specimens cited in 
1938. Those from the Guianas should be rechecked, since both T. 
macrophylla and T. yuyanensis grow there. 

Specimens of T. guyanensis ex.\mined: 

*In the "Index Filicum'^ the citation for this combination is given a^ C. 
Chr. Bot. Tidsskr. 26(2): 285. 1904, but at this place it is given as merely 
Leptochilus fendleri (Eat.), with no basionym cited. There is, in fact, no Eaton 
name that could be a basionym. As shown in the "Index Filicum,*' Chnstensen 
was under the impression that the name Meniscium fendleri Eaton existed 
and that Acrostichum fendleri Baker was a transfer of it to Acrostichum, but 
Eaton never published such a name at the place cited by Chnstensen or 
elsewhere, as pointed out by Maxon (Contr. U.S. Nat. Herb. 24: 62. 1922, 
footnote), and Baker's A. fendleri was a new species. However, Chnstensen s 
reference to Acrostichum fendleri Baker under Leptochilus fendleri in the Index 
Filicum*' serves to validate the combination, with the change of parenthetical 
author from Eaton to Baker, 

20 American Fern Journal 

Trinidad: Orange Road, Homberslcy 226 (US). Brazil, Briti(m, Briiton & 
Freeman 2121 (US); Without locality, Fendler 88 (NY, US); Bot. Card. Herb. 
1584 (US). 



Maratakka-Saparra, Gonggn'jp & Stahel 971 (U, US). Bakhuis Mountains, 
between Kabalebo River and Left Coppename River, Florschuiz & Maas 

305 4 (U). 

British Guiana: Barina River, Jmman (NY, 8 sheets). Demerara River, 
Jenman (NY). . . 

These two species, T. macrophylla and T. guyanensis, are most 

likely to be confused with T. chrysodioides (F^e) ]\Iorton and its 

variety goyazensis, which also have the fertile leaves smaller than 

the sterile, the blades being therefore subdimorphic. The other 

large species thd, might be confused are included in the following 

Sporangia non-setose. 

Sporangia confined to the arcuate cross-veinlets; bases of pinnae broad, 

not cuneate. 

L ■ 

Pinnae broadest at base (Florida, West Indies, Margarita Island 

[Venezuela]) T. reticulata (L.) Proctor 

Pinnae broadest above base (Costa Rica). 

T. turrialbae (Rosenst.) Morton 
Sporangia on the arcuate veinlets, the excurrent veinlets, and some- 
times also on leaf tissue; pinnae cuneate at base (Brazil, Peru, 
Guianas, Amazonian Venezuela). 

_ T. macrophylla (Kunze) Morton 

Sporangia setose on the stalk or annulus. 

Sporangia setose on the annulus;' arcuate veinlets nearly straight, the 

excurrent veinlet often reduced to a mere knob (Trinidad, 

Tobago, Guianas) T. guyanensis (Fee) Morton 

Sporangia setose only on. the sporangium stalk ;' excurrent veinlet well 

developed, not bearing sporangia (Guianas, Brazil). 

T. chrysodioides (Fde) Morton var. chrysodioides* 
T. c. var. goyazensis (Maxon & Morton) Morton 



theTetnP^rpS i^ sometmies not easily determinable. It is likely that 

^ThP ,^;f' k deciduous and sometimes not present on mature material. 

Tian^r Kn^Tt^v." '^^''y^'^^^oides IS accepted here as construed in our 1938 

type is locat^ correctly or not is not really determinable until F^e's 

Thelypteris normalis Complex 21 

helypteris normalis Complex 
Southeastern United States^ 

A. Reid Smith 

The species of the Thelypteris normalis complex in the south- 
eastern United States and the West Indies have long been acknow- 
ledged as taxonomically troublesome. The first person to mono- 
graph the group was Christensen (1913), who treated it under 
Dryopteris, subg. Cyclosorus, More recent workers have also 
recognized the integrity of the group, but generally have treated 
it as a subgroup of Thelypteris (Small, 1938; Wherry, 1943, 1964). 

The chief problems encountered by Christensen and others 
have been the extreme variability in the species and the large 
number of '^forms'' which are seemingly intermediate between 
species. Robert and Edward St. John, avid collectors of Florida 
ferns in the 1930's, added to the proliferation of names when 
they described numerous Florida ^^endemics/' The assiduousness 
of previous workers, however, has laid the groundwork for more 
detailed studies utilizing cytology and field-to-greenhouse trans- 
plants, an approach which I believe has clarified the systematics 
of the group. 



A revision of Thelypteris sect. Cyclosorus in the New World is 
currently being prepared for future publication. However, in 
order that the important conclusions of this study might be more 
immediately available to those with an interest in ferns of the 
southeastern United States and to those involved in the prepara- 
tion of floras, it seems appropriate to publish a portion of the study 
dealing with the species of this area. Following a key to the species 
and varieties is a synopsis of the southeastern United States 
taxa, excluding T. totta (= T. gongylodes) , which is only rather 
distantly related to species in the T, normalis complex. Important 
aspects of synonymy, habitat, distribution, and a discussion of 

^ Much of this research was conducted under a National Defense Education 
Act Fellowship. 

22 American Fern Journal 

the salient differences among the taxa are given, together with a 
tabic summarizing the cytological findings of this study. Docu- 
mentation of chromosome numbers will be presented elsewhere. 

Key to the Species and Varieties of Thelypteris sect. 
Cyclosorus in the Southeastern United States 

1 . Basal veins of adjacent segments united below the sinus with an excurrent 

vein leading toward the sinus; costules, veins, and often lamina 
above hairy. 
2. Rhizomes long-creeping, black, nearly naked; costae below with light- 
colored scales; lamina, veins, costules, and costae above glabrous, 
or sparsely pubescent with very short hairs. 
3. Lamina and veins pubescent below; indusia and costal scales 

hairy T. totta var. hirsufa 

3. Lamina and veins glabrous below; indusia and costal scales 

glabrous or with sparse hairs T, iotta var. totta 

2. Rhizomes short-creeping or suberect; costae below without scales; 

adaxial surface of lamina glabrous or hairy; veins, costules and 
costae above moderately to rather densely hairy with rather 
long stout hairs. 

4, Costae below with predominantly short hairs uniform in length 

(less than 0.2 mm and usually less than 0.1 mm long);- 
excurrent veins mostly greater than 2 mm long; stipes 
purplish; fronds with usually more than two pairs of greatly 

reduced pinnae at the base 1. T. dentala 

4. Costae below with most hairs greater than 0.3 mm long (some ex- 
ceeding 0.5 mm), not uniform; excurrent veins less than 2 
mm long; stipes stramineous; fronds with 0-2 pair(s) of 
shghtly reduced pinnae at the base. 

2. T. quadrangularis var. versicolor 
1. Basal veins of adjacent segments free below, or connivent at the sinus; 

costules, veins, and lamina ahove with or without hairs. 
5. Rhizomes suberect to erect, massive; basal segments of the lower and 

medial pinnae usually narrowly elongate parallel to the rachis; 
auricles (enlarged l:)asal acroscopic pinna segments) often pre- 
sent at the base of the lower pinnae; costae below without 

scales 3^ y ^ patens var. patens 

5. Rhizomes long-creeping to short-creeping, the stipes arcuate at the 

base; basal segments of the lower and medial pinnae either 
reduced, the same size as, or slightly longer than more distal 
pinnules; auricles small or absent; costae below with or without 
small scales. 

Thelypteris norma lis COMrLEX 23 

6. Costae, costules, and veins above with at least a few rather stout 


hairs mostly greater than 0.3 mm long; lamina above 
often rather glandular; scales absent on mature fronds on 
rachis and costae below. 
7. One or two pairs of basal pinnae somewhat reduced; rhizomes 

short-creeping, sometimes appearing suberect; vena- 
tion variable, even on the same frond, from anasto- 
mosing with a short excurrent vein to connivent at 
the sinus; lamina above often somewhat hairy; veins 
above always with stout hairs, many greater than 
0.4 mm long 2. T, qiiadrangularis var. versicolor 

7, Lowermost pinnae usually not reduced; rhizomes short- 

creeping to long-creeping; veins connivent at the 

or the distal one of each pair meeting the margin 

slightly above the sinus; lamina above glabrous or 
sparsely hairy; veins above with or without long 

stout hairs •. . .4. T. norinalis 

6. Costae, costules, and vems above glabrous or with very thin 

short hairs mostly less than 0.2 mm long; lamina above 
eglandular; a few scales persistent on rachis and sometimes 
costae below (absent in T. ovata var. ovata). 

8. Terminal "pinna" usually at least 5 times as long as wide; 

scales on costae below often numerous; lamina 
coriaceous; lower pinnae (at their widest point) often 
less than 1.2 cm wide, incised 0.5-0.7 of their width. 

5. T. migescens 

8. Terminal "pinna" less than 5 times as long as wide; scales 

on costae below absent or relatively sparse; lamina 

chartaceous to subcoriaceous; lower pinnae (at their 

widest point) usually greater than L2 cm wide, incised 

• 0.6 to often more than 0.8 of their width. 

9, Scales usually absent on the costae below; lamina above 

jrlabrous . . . ." 6. T, ovata var. ovala 

9. A few very narrow scales usually present on the costae 

below; lamina above minutely pubescent or 
glabrous 6a. T. ovala var. lindheimeri 

1. Thelypteris dentata (Forssk.) E. St. John, Amer. Fern J 

26:44. 1936, 



Yemen, Forsskal s.n. (C-Herb 

24 American Fern Journal 

Pohjpodium molle Jacq. Coll. 3: 188. 1789, non Schreb. 1771; nee AIL, 
1785. Holotype: cultivated in the Botanical Garden, Schoenbrunn, and 
said (probably incorrectly) to originate from Caracas, Venezuela (W! 

photograph UCI). 
Aspidium molle Swartz, J. Bot. Schrad. 1800(2): 34, 1802. To be con- 
sidered a new name under Art. 72 (Note) for Pohjpodium molle Jacq., 


Aspidium violascens Link, Hort. Berol. 2: 115. 1833. Holotype: cultivated 
in the Botanical Garden, Berlin, from plants said (probably incorrectly) 
to originate from Brazil (B, not seen; isotypes HBG! photographs UC! 

US!; LI photograph UC! USI). 

Dryopteris mollis (Swartz) Hieron, Hedwigia 46: 348. 1907- 

Dryopteris denlata (Forssk.) C. Chr., Dansk. Vid. Selsk. Skr. VIII, 6: 24. 

Thelypieris reducia Small, Ferns Southeastern States. 254. 1938. Lectotype 
(chosen herein): "Inter Bowling Green et Fort Green [Hardee Co.], 
Florida", 7-11 Dec. 1934, E. SL John, R. SL John, W. A. Knight, and 
/. K. Small, s.n. (NY!). 

Habitat and DisTRiBrTioN: RoadsideSj pastures, edges of 
woods, and other disturbed habitats; also limestone sinks. Tropical 
and subtropical North and South America, with the possible 
exception of the Andean countries. This species is a common 


weed" of waste areas in tropical Africa, Asia, and the Pacific 
Islands. In Florida, T. dentata is probably confined to hammocks, 
especially those of a limestone nature. 

There is the strong indication that this species is not native to 
the New World but has been introduced rather recently (Smith 
in Strother and Smith, 1971). The earliest United States collection 
I have seen is from near ]Mobile, Alabama, in 1904. It was probably 
not collected in Florida until 1930, when it was found in Her- 
nando County. .Subsequently, it has been found in at least ten 
Florida counties and appears to be increasing its range, both in 
Florida and throughout the New World. Single collections I have 
seen from Georgia, Kentucky, and Louisiana are probably all 
escapes from cultivation. That T, dentata has escaped from culti- 
vation is not surprising, since it is a very common fern in green- 
houses in the United States. Its native relative, T. quadrangularis, 
is rarely cultivated- 

Thelypteris normalis Complex 25 

From T. quadrangularis, T. dentala differs in the longer 
excurrent vein, the uniformly short pubescence below (long or 
long and short hairs mixed in T. quadrangularis) , and the purplish 
color of the stipe and rachis (stramineous in T, quadrangularis) . 
Thelypteris dentata has been counted many times as tetraploid 
(n = 72) from both Old and New World collections, while T. 
quadrangularis is diploid. 

2. Thelypteris quadrangularis (Fee) Schelpe var. versicolor 

(R. St. John) A. Reid Smith, comb. nov. 

Thelypteris macilenta E. St. John, Amer. Fern J. 26: 50, 52, pL 5. 1936. 

Holotype: 7 mi NW of Brooksville, Annuttalagga Hammock, Florida, 

E. St. John W4 (NY! photograph UC!; isotypes MO!, 2-NY!, US!). 
Thehjpteria versicolor R. St. John in Small, Ferns Southeastern States, 250, 

cu7n tab. 1938. Holotype: Brooksville, Hernando Co., Florida, ff. SL 

John 109 (NY! photograph UC!). 
Thelypteris X versicolor R. St. John {pro sp.). StaL emend.' in Wherry, 

Southern Fern Guide 120. 1964. 

IL\BiTAT AND DisTEiBUTiox: Woods and limestone sinks along 
the southeastern coastal plain from South Carolina to eastern 
Texas; also known from Cuba. Other varieties of T. quadrangularis 
are found in Mexico, Central and South America, Africa, and 

probably Malesia (Holttum, pers. conim.)- 

Although previously considered either a species or a hybrid 
between T. dentala and T. normalis, T. versicolor is best treated 
as a variety of the widespread T. quadramjvlaris, differing from 
the type of T. quadrangularis (from French Guiana) by its conni- 
vent or weakly united veins, short-creeping rhizome, and distri- 
bution. Typical T. quadranyularis from the Lesser Antilles, and 
Central and South America has united basal veins and an erect 
rhizome. In the West Indies, var. versicolor grades into two other 
varieties of T. quadrangularis, including the type variety, making 
specific recognition impossible. 

Wherry (1964, p. 120) treated T. versicolor as a hybrid, stating 
that the sori tend to be "sparse and poorly developed, and [the] 

26 American Fern Journal 

spores mostly aborted. [Thelypteris versicolor is] Manifestly of 
hybrid origin between distantly related taxa/^ meaning T. dentata 
and T, normalis, I have examined spores of versicolor throughout 
its range and find that they are normal in appearance. Chromosome 
counts from Louisiana and Florida (Table I) show that var. 
versicolor is a diploid with 3G pairs of chromosomes at meiosis. 
This contrasts with the tetraploid condition in both T. normalis 
and T. dentata. 


Thelypteris macilenta, km 
specimens propagated from the original clone, appears distinct 
biit is no more than an aberrant element within var. versicolor 
The rhizome habit, small size of the fronds, reduced pinnae below, 
pinna shape, and stout hairs on the ultimate veins and costules 
both above and below all suggest this relationship. 

3. Thelypteris patens (Swartz) Small, Ferns Southeastern 

States 243. 1938. 

Polypodium patens Swartz, Prodr. 133. 1788. Lectotype (chosen herein): 
Jamaica, Swartz s.n. (S-PA! photograph UC!). 

Aspidium stipulare WDld., Sp. PI. ed. 4, 5: 239. 1810. Holotype: Plumier, 
Tract. Fil. pi 23 ! 

Aspidium macrourum Kaulf. Flora 6: 365. 1823. Holotype: Martinique, 
Sieher 354 (not seen; isotypes L, photograph US!, M!, MO!). 

Nephrodium albescens Desv. Mem. Soc. Linn., Paris 6 : 258. 1827. Holotype: 
Jamaica, collector unknown (P, not seen, photograph US!). 

L ■ 

Habitat and Distribution: Common along roadsides and 

paths, edges of woods, and open woods in the West Indies, southern 

Mexico, Central America, and south to southern Brazil and Bolivia. 

From the United States I have seen only a single specimen: 

Florida, Dade Co., S of Aliami, Feb.-IMar. 1905 A. A. Eaton 
s.n., (US!). 

Thelypteris patens var. patens can be distinguished from most 
JSew \\ orld taxa in Cyclosorus by its erect rhizome and by the 
stipe base scales, which are ovate-lanceolate, light brown, and 
glabrous. The basal pinna segments in var. patens are either 
elongate and acuminate, or the acroscopic segment is expanded 

Thelypteris noraialis Complex 27 

into a deeply pinnatifid auricle up to 5 cm long. Without the 
rhizome, a specimen of T, patens is sometimes difficult to distinguish 
from specimens of T. normalis. The latter species has hairs on the 
eostules and ultimate veins above and the lamina above is fre- 
quently glandular. The blade "above in T. patens var, patens is 
glabrous except for hairs along the costa. All specimens of 7". patens 
thus far counted have been tetraploid (n = 72), 

4. Thelypteris normalis (C. Chr.) Moxley, Bull. So. Calif. 

Acad. 19:57. 1920. 

Dryopieris patens var. gJanduhsa A. A. Eaton, fiull. Torr. Chib 33: 477. 

1906. Holotype: ColwelFs Hammock, Dade Co., Florida, A. A. Eaton 

185 (GH! photographs MO!, UC!). 
Dryopteris normalis C. Chr.. Ark. for Bot. 9(11): 31. 1910. Syntypes: 

Jamaica, Jenman Herb, s.n. (NY!). 
Thelypteris saxatilis R. St. John in Small, Ferns Soutlieastem States 

236. 1938. Holotype: Ellaville, along the Suwanee River, [Madison Co.], 

Florida, R, SL John 86 (NY!). 

Thelypteris unca R. St. John in Small, Ferns Southeastern States. 246. 
1938. Holotype: Costello Hammock, Dade Co., Florida, R. St. John 172, 
2 sheets (NY! photograph UC!). 

Thelypteris macrorhizoma E. St. John, Amer. Fern J. 32: 146. 1943, Holo- 
type: Sheep Island, The Cove, 6 mi from Floral City, Citrus Co., Florida, 
E. SL John 1273 (FLAS! photograph UC!; isotypes OH?, US! photograph 


Habitat and Distribution: Common in woodlands, limestone 
sinks, gullies, and wet roadsides in open or partially shaded situ- 
ations. United States: coastal plain from South Carolina to 
eastern Texas. Bermuda, Eahama Islands, Greater Antilles, 
Tortola, and from Costa Rica north through the Yucatan peninsula 
to southern Tamaulipas, ]\Iexico. Rare in the Lesser Antilles, 
northern Venezuela, and northeastern Brazil. Reports by Morton 
(1967) that it occurs south to Bolivia are dubious. 

Thelypteris normalis is a well-characterized species but it has 
been either too broadly or too narrowly construed by most 
taxonomists. It can best be distinguished from T. ovata by the 
usually hairy eostules and veins above, and by the stipitate glands 

28 American Fern Journal 

on the adaxial surface of the lamina. In general, the pinnae of T. 
normalis are not so deeply incised as those of T, ovata and the 
segments are less obhque, Sori in T. normalis are medial to supra- 
medial, whereas in T. ovata they are supramedial to submarginal. 
The presence of single-celled glands on the sporangial stalks and 
the absence or paucity of glands on the indusial margin are excel- 
lent characters for distinguishing T. normalis from T. ovata, where 

r ^^ 

the situation is reversed (glands always absent on sporangial 
stalks but present and often abundant on the indusial margin). 
Throughout most of its range, T. normalis is easily distinguished 
from T, quadramjnlaris by venation, but in the southeastern 
United States, T. quadrangnlari s var. versicclor often has the basal 
pair of veins only weakly united or merely connivent at the sinus. 
Thelypteris normalis can then best be identified by the lowermost 
pinnae being usually not reduced (the lamina deltoid rather than 
ovate-lanceolate in outline) , by the lack of hairs on the leaf tissue 
between the veins above, by its generally longer creeping rhizome, 
and by the slightly thicker blade texture. 

Thelypteris normalis hybridizes with T. augescens, T. ovata var. 
ovata, and probably T. quadrangulariSy although hybridization 
with the last-named taxon has not yet been cytologically verified. 
In all cases examined the resulting hybrids are sterile. 

Morton (1967) attempted to show that there .are at least two 
names with priority over Dryopteris normalU C. Chr. One of 
these, Aspidiiim germanii Fee, I have placed in synonymy under 
Thelypteris quadrangularis. Unfortunately, the type of the second, 
Nephrodium kunthii Desv,, has not been available for study, but 
I believ6 that it very likely is not T. normalisj but rather a variety 
of T. patens. 

Thelypteris saxatilis and T. unca are both easily included witliin 
the latitude of variation of T. normalis. Thehjiteris saxatilis is a 
somewhat smaller form than is usual for the species ; some of the 
specimens so labeled by St. John are referable instead to T. ovala 
rather than T. normalis. Thelypteris unca differs from most speci- 
mens of T. normalis mainly in its slightly more falcate pinnules. 
Thelypteris macrorhizoma is less pubescent than "typical" normalis, 

Thelypteris normalis Complex 29 

but differs in no other significant way. 


5. Thelypteris augescens (Link) Munz & Johnston, Amer 

Fern J. 12: 75. 1922. 

Aspidiumaugescens Link, Fil. Sp. 103. 1841. Type. Cuba, Otto 89, right hand 
plant (B! photograph UC!; additional authentic material is at BR! 
and UC!). Cultivated in the Botanical Garden, Berlin, from plants said from Caracas. Christensen (1913) discussed the origin of the type 
material and concluded, no doubt correctly, that the type locality must 
have been Cuba. 

Habitat and Distribution: Dade Co., Florida, Cuba, and 

on the islands of Andros and New Providence in the Bahamas. 
Generally on exposed limestone outcrops in moist to moderately 
dry open sites. 

. Thelypteris auijescens can be distinguished readily from its close 
relative T, ovata by the more numerous paleae on the costae 
below, by the more coriaceous blade, by the greater pinna length/ 
width ratio, by the less deeply incised pinnae, and by the usually 
distinct terminal ''pinna/' Glands are absent on the sporangial 
stalks, but usually present on the indusial margin, as in T, ovata. 
The West Indian T, sena greatly resembles T. atigescens but 


costal scales 

and more distinct terminal ''pinna.'' Additionally, the basal 
segments of the lower pinnae in T. aWjesceus are never reduced as 
is usually the case in T. serra, but are as long as or sHghtly longer 
than the more distal piimules. Reports of T. sevra in Florida 
(Small, 1938) are the result of confusion with T. augescens. From 
the Alexican and Central American T. jmhervh, T, augescens 
differs in its more coriaceous texture, the longer and narrower 
lateral pinnae, and the usually distinct terminal ''puma.'' The 
costal scales of T. augescens are longer, more numerous, and of 
lighter color than those of T. puherula. The geographic ranges of 
T, augescens and T, puherula do not overlap. 

Thelypteris augescens hybridizes with T. normalis in Cuba and 
Dade Co., Florida. Hvbrids also occur with T. ovata var. ovata in 

30 American Fern Journal 


are sterile and the spores abortive. 
6. Thelypteris ovata R. St. John in Small, Ferns Southeastern 

States 239, cum tab. 1938. 

Dryopieris normalis var. harperi C. Chr. Dansk. Vid. Selsk. Skr. VII. 
10: 182. 1913. Holotype: Hawkinsville, Pulaski Co., Georgia, R. M. 
Harper 1382 (US! photograph UC!; isotypes MO! NY! photograph UC!). 

Holotype: Lecanto, Citrus Co., Florida, R. St. John 394 (NY!).. 

Habitat and Distribution. Hammocks and limestone sinks 
along the southeastern coastal plain from S. Carolina to Alabama 
Andros and New Providence in the Bahamas. 

6a Thelypteris ovata var. Undheimeri (C. Chr.) A. Reid Smith, 

comb. nov. 

Dryopieris normalis var. Lindheimer C. Chr. Dansk. Vid. Selsk. Skr. VII, 

w: inz. iyi3. Holotype: Texas, F. Lindhe 
UC ! ; isotypes MO ! UC ! US ! photograph UC !) . 
lelypteris X Undheimeri (C. Chr.) Wherrv. S( 

xiABUAT AND Distribution: Riverbanks and moist canyons 
from central Texas south to northern Veracruz and Puebla; 
apparently most common in Texas and the states of Nuevo 
Leon, Tamaulipas, and San Luis Potosi in Mexico. 

The characters which best distinguish T. ovata from its closest 

relatives, namely T. normalis and T. aucesceris, have already been 

Two forms of T. ovata var. ovata occur in southern Florida which 
are rather different from each other, and at the same time are 
unlike specimens from central and northern Florida. One of these 
^ found chiefly in the Everglades and is characterized by fronds 
frequently greater than one meter long and pinnae 18-25 cm long. 
The second form occurs in lime sinks and along roadsides and 
nver banks. Its fronds are seldom greater than 0.5 m long and its 

Thelvpteris normalis Complex 31 

pinnae are much shorter^ often less than 10 cm long. These two 
forms are alike in nearly all other characters. When grown under 
uniform greenhouse conditions, these forms show much less pro- 
nounced size differences and there is little doubt that they belong 
to the same species. 

The presence of linear scales (mostly 0.5-1.0 mm long) on the 
costae below is the best character for distinguishing var. lind- 
heimeri from the type variety. In addition, the blade is usually 
less broadly deltoid than in var. ovala. There are often minute 
hairs about 0.1 mm long on the lamina above in var. lindheimeri] 
these are absent in var. ovata. The two varieties are allopatric. 

Table I. Summary of chromosome numbers in species and hybrids of 
Thelypteris sect. Cyclosorus from the southeastern United States 
and neighboring regions,^ 


Chromosomes 'populations 

Taxon at meiosis sampled^ 

T. augescens 72 II " 2 

T. dentata 72 II 3 

T, normalis 72 II 28 

T. ovata var. ovata 36 II 13 

T. patens 72 II 1 
T, quadrangularis var. 

versicolor " 36 II 9 


T. augescens X normalis ca. 36 II + 72 I 2 

T. augescens X ovata ca. 36 II + 36 I 2 

T, normalis X ovata ca. 36 II + 36 I 4 

^ For most populations, several plants were counted 

Literature Cited 

Christensen, C. 1913. A monograph of the genus Dryopteris. Part I. The 

tropical American pinnatifid-bipinnatifid species. Dansk. Yid. Selsk. 
Skr., Ser. VII. Nat. og Math. Afd. 10: 55-282. 

Morton, C. V. 1967. Studies of fern types, I. Contr. U. S. Nat. Herb. ^Si 




32 American Fern Journal 


Strother, J, L., and A. R. Smith. 1970. Chorology, collection dates, and 

taxonomic responsibility. Taxon 19: 871-874. 
Wherry, E. T. 1943. Observations on Florida ferns. Amer. Fern J. 32 : 139-145. 
. 1964. The Southern Fern Guide. Southeastern and South-Midland 

United States. Doubleday, Garden City, New York. 

Herbarium, Department of Botany, University of Cali- 
fornia, Berkeley, California 94720. 

Asplenium pinnatifidiun X trichomanes^ANew 

Record for Indiana 

Gerald J. Gastony 

One of the most interesting aspects of the 1970 foray of the 
American Fern Society was the discovery of a new record for the 
state of Indiana, the rare hybrid Asplenium pinnatifidum X 



this natural hybrid, which was previously known only as a single 
plant from southern Illinois (Wagner & Wagner, 1969). The 
specimen here reported was discovered by the author — and nearly 

in a pocket on a south-facing 


sandstone cUff at McBride's Bluff along the east fork of White 
River in Martin county, Indiana, about six miles north of Shoals 
on August 22, 1970. Rolla Tryon and the author had been attracted 
to this particular portion of the area's extensive sandstone cliffs by 
Amy and James :\Iason and Lynda Cole, other foray members, 
who were investigating a peculiar variant of Woodsia ohtusa on an 
adjacent east-facing spur of the outcrop. 

Like the specimen reported by Wagner and Wagner, the present 
hybrid is morphologically intermediate between the parental 
species. The pinnae are quite like those of A. trichomanes {Plate 
6), but express the influence of the A. pinnatifidum genome in 
being more broadly attached to the rachis. The influence of A. 
pinnatifidum is also evidenced by the long-tapering lobed apex of 
several of the hybrid's leaves. The petiole and proximal part of 

American Fern Journal 

Volume 71, Plate 6 


""^■•yitifi^^ mV'.-.«.«.'.-.*.'».'.-'.".-'.'WW.^ -In'r'V ' 


"■' ¥>:-, 

•MSW«SNX«*»*>^,,.;^^ ,.^;^ 


2 mB» 




<P "-"■^-^"-^-"-V-''-Vf-"- '-^-^ -TJ^'^^*- " ' - -- - 



near Shoals, Indl\na. Fig. L Representative leaf of A. pinnatifidum 
NuTT. Fig. 2. Representative leaf of A. trichomanes L. Fig. 3. A. pin- 
natifidum X trichomanes. 

34 American Fern Journal 


the racliis is brown (with brown streaks often extending to the 
second to fourth pair of pinnae) and half-terete to adaxially chan- 



minute, clathrate, trichomoid scales on the petiole and rachis of 



of the hybrid are clearly abortive. Occasional sporangia in several 
sori are rather normally developed, but even these evidently bear 
aborted spores whose number is uncertain but appears to be con- 
siderably fewer than the 64 of the parental species. This may 
indicate the incipient development of a sporogenetic system co- 
ordinate with apogamous re})roduction. 

Because fixatives were unavailable in the field and the specimen 
did not appear fresh enough for cultivation upon arrival at Indiana 
University, the hybrid was not investigated cytologically. Nor have 
the populations of ^ . trichornanes at :\IcBride's Bluff been examined 
to determine which of the ploidal levels known for this species may 
be represented. The leaves of the present specimen are morpho- 
logically quite similar to those figured by Wagner and Wagner in 
which the parental A. trichomanes population was diploid. Thus it 
IS not morphologically apparent that A. trichomanes of higher 

ploidy than diploid has been involved in the parentage of this 
hybrid . 

Voucher material of this specimen, Gastony 949, has been 
deposited in the Indiana University Herbarium at Bloomington 
(IND) and at the Gray Herbarium (GH). 

Literature Cited 

Wagner WH. Jr. 1954. Reticulate evolution in the Appalachian Aspleniums. 

Evolutions: 103-118. . ^^ ^ 

Wagner W. H. Jr. and F. S. Wagner. 1960. A new natural hybrid in the 

Appalachian Asplenium complex and its taxonomic significance. 

Brittonia 21 : 178-186. 

Department op Botany, Indiana University, Bloomington, 
Indiana 47401. 

DwARD Ostrich Ferx 35 

A Dwarf Ostrich Fern 

H. Lou Gibson 


In mid-sunimer 1956^ I found a small, isolated, patch of ferns 
of a species that I did not recognize. The ferns were growing in 
a strip of moist, alluvial woods along the bank of the Moose River 
m the Adirondacks, near McKeever, Herkimer County, New 
York. They appeared similar to typical Matteuccia struthiopferis 
var. pensylvanica (Fig, 1), but had slightly more rounded and 
much shorter fronds {Fig, 2), They did not appear to be repro- 
ducing by offsets from their rhizomes, nor do I recall seeing any 
fertile fronds. The patch was separated by about 50 feet from 
Ostrich Ferns growing along the river bank. The land there was 
about one foot higher and somewhat drier than in the rather 
mucky flat where the small Ostrich Ferns grew. 

I brought a few plants back to my wildflower garden. They 
persisted successfully and did not change in character with re- 
spect to size, offsetting, or fertihty. In 1967 one plant did produce 
a stunted and twisted ''fertile'' frond about one inch long, tight 
to the crown. But no prothallia could be grown from it. 

In 1964 I studied stem sections in an attempt to identify the 
species. The vascular strands of the dwarf Ostrich Ferns seemed 
identical to those of the regular ones. 


In order to exclude the possibility that soil conditions mmy 
garden maintained an environmental dwarfing, I did a few ex- 
periments. Small plants were paired as to size with young offsets 
from large Ostrich Ferns. Three pairs were planted in various 
parts of the garden. In each case the regular ferns grew to a large 



the opinion that the dwarf condition is genetically fixed. The 
dwarf plants grew equally well in wet and drier areas of the 
garden, whereas the normal plants attained greater heights in a 
wet section than in a dry one. 
In 196o I returned to the natural site to study the original 

American Fern Journal 

Volume 71, Plate 7 

Matteuccia struthiopteris fronds. Fig. 1. Normal frond 60 inches 
LONG. Fig. 2. Dwarf frond 15 inches long. 

Shorter Notes 37 

patch. Trees had been cleared along the strip for road buildmg, 
causing a denser undergrowth. A washed-out dam had lowered 
the river by about two feet. Ostrich and Cinnamon Ferns were 
more profuse than formerly. The area was searched, but every 
likely Matteuccia of small stature could be traced by the long- 
creeping rhizomes back to a large, full-grown Ostrich Fern. A more 
detailed and lengthy search, perhaps with a ready means for 
identification, might rediscover the original dwarf population. The 
small ferns are probably overgrown by the normal species. 

Some perusal of the literature and consultations with fern 
authorities have not uncovered any mention of a dwarf variety of 
the Ostrich Fern. Perhaps this report will prompt further 

Rochester Academy of Science, Rochester, N. Y. 14617, 

Shorter Notes 


In Copeland's ''Genera Filicum," Ntphidium, based on .V. arneri- 
mnum (Hook.) J. Smith (Hist. Fil. 99. 1875), is recognized as a 
distinct; monotypic genus, a native of the Andes of Ecuador. This 
species has been generally placed in Cydophonis (now correctly 
Pyrrosia), which is a genus strictly confined to the Old World. 



closely allied. As Copeland pointed out, Pyrrosia has stellately 
branched peltate scales (Fuj. S) on the undersurface, Avhereas 
Niphidinm has (under the compound microscope) beautifully 
clathrate scales bordered with elongate, hyaline hairs {Fig. 1). 
The hairs are usually forked at the apex and two cells wide at the 
base (Fig, 2) ; each cell forms one of the forks. These hairs are so 
numerous and dense as to make the lower surface appear densely 
tomentose. But the frond scales of Xiphidhim are not truly peltate 
and stellate, and so a resemblance to Pyrroma is strictly super- 
ficial. In its Anaxetum-\\ke venation and its large, round sori 

Amekican Fekn Journal 

Volume 71, Plate 8 


Shorter Notes 39 

uniseriate between the prominent lateral veins Niphidium is like 
Pessopteris, and may be related to that, which is however essentially 
scaleless on the lower surface. 

In calling this plant Niphidium americanmn (Hook.) J. Smith, 
Copeland overlooked an earlier specific name that is available. 
In his paper on Cavanillcs' fern types, Christensen indicated that 
Poly podium longijolium Cav., from the type specimen, is identical 
with N, americanum. Therefore, the following new combination 
is necessary: 

Niphidium longifolium (Cav.) ]Morton & Lellinger,'comb. nov. 

Polypodium longifolhtm Cav. Descr. 245. 1802. Type: Pelileo, near Mount 

Tiinguragua, Ecuador, Nee (MA, seen by Christensen). 

Polypodium americanum Hook. Sp. Fil. 5: 54. 1864. Syntypes: Cuenca, 

Ecuador, Jameson, and Banos, Ecuador, Spruce 524^ (both pre- 
sumably K). 

Niphodolus americamis (Hook.) Diels in Engl. & Prantl, Nat. Pflanzenfam. 

1(4): 325. 1899. 
• Cyclophones americamis (Hook.) C. Chr. Ind Fil. 198. 1905. 

Cyclopkorus longifoliiis (Cav.) C. Chr. Dansk Bot. Ark. 9(3): 11. 1937. 

It should be pointed out that under Pyrrosia the epithet longi- 
folia is not available because of the quite different Old World 
species P. longijolia (Burm. f.) ]\Iorton (J. Washington Acad. Sci. 
36: 168. 1946).— C. V. ^Morton and David B. Lellinger, 
National Museum of Natural History, Washington^ D, C 20560, 

Bulbous Adder's-tongue Commox in Louisiana. — Bulbous 

Adder's-tongue is a very small, easily overlooked fern. I first saw 
it in IMarch, 1970, accidently while looking for Oenothera spachiana 
seedlings for my wildflower garden. Most manuals consider this 
fern to be rare or at least uncommon. Brown and Correll (Fern 
and Fern Allies of Louisiana, 1942) listed it from four southeast 
parishes (East Feliciana, East Baton Rouge, Livingston, and 
Orleans), one southwest parish (Jefferson Davis) and two northeast 

nnris^pc r"Rir>lilQni1 nj^A PoUItvpU^ Thpv notpd that it waS common 

Fig. 1. Scale from the ab.vxtal surface of Niphidium longifolium. 

X 24 (GiLER o9). Fig. 2. Same, detail of base of double-celled hair 


stichoides, X .50 (Holxtum 18054). 

40 American Ferx Journal 

only on the Copenhagen Prairie of Caldwell Parish. The University 
of Southwestern Louisiana has specimens from Bienville, Winn, 
Caldwell, and Rapides Parishes. Louisiana Technological Univer- 
sity has a specimen from Union Parish. The small number of 
specimens and the information on the labels indicate that most 
collectors consider this fern to be rare. 

From mid-?^Iarch to mid-]May, 1970, I made field trips looking 
specifically for Ophioglossum crotaloph oroides. I have collected the 
fern in the following twenty-six pai-ishes: Allen, Beauregard, 
Bienville, Bossier, Caddo, Caldwell, Catahoula, Claiborne, DeSoto, 
Evangeline, Franklin, Grant, Jackson, LaSalle, Lincoln, INIore- 
house, Natchitoches, Ouachita, Red River, Richland, Sabine, 
Union, Vernon, Webster, West Carroll, and Winn. Other parishes 
could have been added had I not been restricted in my collecting 
by a full teaching load. The specimens referred to above raise the 
number of parishes where this fern is known to grow to thirty-two. 

Ophioglossum crotalophoroides is apparently absent from four 
parishes I checked: East Carroll, :\radison, Tensas, and Concordia. 
They lie along the Mississippi River, and I could not find any 
sandy soil in them. No plants were found on the clay soils character- 
istic of these parishes. All collections were from sandy soils of 
cemetaries, pastures, road banks, power lines, pipe lines, etc. 
Typically these sites are mowed during the summer. In such 
habitats the plants of Bulbous Adder's-tongue are extremely nu- 
merous. Some cemetaries I visited contained thousands of these 
plants. I think there are almost as many (or as many) Ophioglossum 
crotalophoroides plants in Louisiana as there are of such common 
ferns as Christmas Fern and Bracken; it sunply has been over- 
looked by collectors. One possible explanation of this is that during 
late March, when 0. crotalophoroides is at its peak, there are no 
common showy associates— except possibly the tiny bluets, Hous- 
tonia pusiUa, H. minima, and H. pygmaea). At this time few 
botanists bother to Collect in such "unfruitful" territory. Also, 
one can walk over those ferns ten times and fail to see them. But 
this small fern can be seen with any degree of regularity only from 
the prone position. I suggest that collectors look specifically in the 

Recent Fern Literature 41 

proper habitats for this fern and that they get down to ground 
level where the plants are. Ophioglossum crotalophoroides is much 
more common than previously supposed. — Dr. R. Dale Thomas, 


71 SOL 

Recent Fern Literature 

Flore de la Nouvelle-Caledonie et Dependances, no. 3, 
Pteridophytes, by G. Brownlie. Published by Museum National 
d'Histoire Naturelle, 16 rue Buffon, Paris V, France, pp. 1-307. 
L I-XXXIX. 1969.— This beautifully prepared book is the first 
fern flora of any of the South Sea Islands for many years. jMany 
of the ferns of New Caledonia are endemic, and extremely inter- 
esting because of their variability, in which they are comparable 
to the ferns of Hawaii, - 

Brownlie's treatment has full keys, descriptions, and citation of 
synonyms, misapplied names, and specimens. The work owes 
much to Madame Tardieu-Blot, who translated the descriptions 
into French and who had the excellent drawings prepared in 
Paris. In fact, the whole style and format is exactly like one of 
^Madame Tardieu's own books on the fern floras of Gabon and 
Cameroun; however, Madame Tardicu disclaims an}'' responsi- 
bility for the taxoiiomic treatment or other details. The comments 
that I give below are on minor matters. They do not detract from 
the value of this excellent book, which should be in the hands of 
all fern students. 

Brownlie splits up the traditional family Polypodiaceae into 13 
families, following Alston (cf. Taxon 5: 23-25. 1956), who had a 
much better treatment in my opinion than that of Ching, Cope- 
land, or other more recent authors. However, one runs into 
difficulties in trying to describe and key out these families, and 
Brownlie's key will not work well for some genera. Grammitidaceae 
has as one of the key characters that the petiole is generally 
articulate, but tliis is never true in this family. Vittariaceae is 


strictly dorsal in Antrophijum. Acrostichum, Syngramma, and 

42 American Fern Journal 

Asplcniopsis, which have the sori strictly dorsal, are keyed solely 
under the heading of sori marginal or submarginal. This merely 
points out that it is impossible to key out the families as recognized 
by single key characters. Probably the only way is to do as Cope- 
land did in his "Genera Filicum," and have Aspidiaceae appear 
in ten different places in the key to the families, Pteridaceae in 
nine places, and so forth. Such keys are essentially keys to genera 
or groups of genera, rather than to families. 

^lany of the family names used are suspect or definitely in- 
correct. Azollaceae is, for instance, attributed to Christensen in 
Verdoorn (1938), but at the place cited there is no Latin diagnosis. 
Presl called his "Lindsaeaceae" a "Sectio," and since a section is a 
group below the rank of a genus, it can not be a proper category 
for a family name. The Code indicates that a family name must be 
called either a family or an "Ordo" (Art. IS, paragraph 3). Vittari- 
aceae was called by Presl a "Tribus," and a tribe is also an in- 
admissible category for a family name. PsUotaceae is attributed 
to Payer (1850), but Payer had it in French as "Psilot^es"; since 
botanical nomenclature must be in Latin, names published origf- 
nally in French are not valid; cf. Code, 1966 edition, p. 213, where 
"Casuarin^es" .Alirbel, 1810, is rejected in favor of Casuarinaceae 
R. Brown, ISU.^ 

Many stated types are in reality lectotypes only (e.g. Divlazium 
echtnatum C. Clir.). Vieillard 1520 from Canala can not be the 
type or lectotype of Blechnum coufusum (Foum.) Brownlie, for 
It was not one of the collections cited; one of the syntypes was 
Vieillard 1520 from Balade. IMany Paris specimens are cited as 
types," I.e. miplicitly holotypes, when they are actually, as in the 
case of Labillardiere species, really isotypes or isosyntypes. The 
type of MtCTolepia is said to be M. speluncae (L.) Moore, but this 
IS impossible, since this was not one of the species originally 
mcluded m the genus. The type of Ladreopsis is Polypodium 
recedens J. Smith, not L. tenera (R. Brown) Tindale, which may be 

citaUoL%fTth4a.'Sp'AW ^^ ^'^f.^^ typographical errors; one noted is the 
ciiaiion ot Athyriaceae Alston as "Ta.xon 5: 36. 1936"; it should be 5: 25. 1956 . 

Hecext Fern Literature 43 

a taxonomic synonym but which is based on an entirely different 
type specimen. 

The only genus in which the treatment is unsatisfying is that 
of Asplenium, in which a number of species and varieties described 
from New Caledonia are placed in a table showing their possible 
origin as hybrids, but with absolutely no discussion of these, or 
any citations, descriptions, or typifications. 

Some of the compound epithets are spelled without a hyphen, 
e.g. Hymenophyllum le ratii and Adiantum capillns veneris, but 
the Code requires such specific names to be hyphenated. The 
specimens that are referred to Lycopodium nutans Brack. (1854) 


are probably L, proUferum Blume (from unpublished annotations 
by Adelbert), and if this is correct the Blume name has priority. 
Cyathea propinqua ]\Iett., a Fijian species of the subg. Sphaero- 
pteriSj is cited as a synonym of C vieiUardii Mett., which belongs 
in subg. Cyalhea, according to Holttum's classification (Blumea 12 : 
272. 1964), Asplenium adiantoides (L.) C. Chr. (1905) is accepted 
as a correct name, but it is an illegitimate later homonym of A . 
adiantoides Lam. (17SG); 1 believe that the correct name is ^- 
polyodon Forst. Thelypieris uligiiwsa (Kunze) Ching has Cheil- 
anthes sefigem Blume (1828) cited as a synonym; T. uliginosa is a 
different species fron the Javan 7". setiyera; the correct name under 
Thelypteris is T, torresiana (Gaud.) Alston, which has recently 
been called Macrothelypferis torresiana (Gaud.) Ching (cf. Holttum, 
Blumea 17: 27. 1969). Marattia rolandi-principis is described as 
being simply pinnate, but it is sometimes bipinnate; Rosenstock 
distributed material as M. rolandi-principis var. subbipinnata 
Rosenst., an unpublished name, and the bipinnate plants were 
collected by Guillaumin and Baumann (no. 8663). Nephrolepis 
floccigera (Blume) ]\Ioore is cited as a synonym of -\^ cordifolia, 
but from the holotype (from Celebes) it is by no means that, for 
it has the pinnae about 12 cm long or more, not about 2 cm long 
as in .V. cordifolia. Blechnum vieiUardii var. simplex Fourn. is a 
nomen nudum at the place cited. 

There are few omissions. The genus Belvisia [Hymenolepis] 
perhaps needs further study; only B. mucronata is reported. 

44 American Fer\ Journal 

Several collections of Hymenolepis revoluta var. planiuscula (Mett.) 
Hieron were cited from New Caledonia by Christenseu (Dansk 
Bot. Ark. 6(3): 59. 1929). Cephalomanes australicnni v. d. Bosch 
was described from the Isle of Pines, which is included within this 
New Caledonian flora. Pyrrosia varia (Kaulf.) Farwell has been 

— L 

reported from New Caledonia but is not accounted for here. 
Otherwise, the only omissions noted are: Blechnum irregulare 
Carr. (New Caledonia, MacGillivray F25, holotype BM, IMorton 
photograph 6685) J S. diversifolium var. paleaceo-seiosum ^osenst^ 
(Fedde Repert. 8: 75. 1910), Lomaria attenuata f. monstrosa 
Compton, L. lenormandii f. apnea Com\)tony A splenium adian- 
toides var. tripinnatum Compton (= A. rohustum Blume?),' 
Cyathea alhijrons var. lata Compton, Hymenophyllum mnioides f. 
amplior Compton, and Lycopodinm sqnarrosum var. pacificum 
Compton.— C.V.M. 

The Ferns aI^d Fern Allies of the Sierra Nevada in 
California and Nevada, by John Thomas Howell. The Four 
Seasons 3(3): 2-18. 1970, — In anticipation of his forthcoming 
*' Flora of the Sierra Nevada," Howell has presented here a list of 
the ferns and fern allies with a key and an indication of the north- 
ernmost and southernmost known stations. There are altogether 
62 species treated, one of which is introduced (Cyrtomium falca- 
turn) and several of which are probable hybrids. "The Four 
Seasons" is a relatively unknown periodical issued quarterly and 
consists chiefly of taxonomic notes on Californian plants. It is 
published by the Regional Parks Botanical Garden, Berkeley, 
Calif. 94708; the subscription price is $6.00 per volume. — C.V.M. 

American Fern Society 

Report of the 1970 Fern Foray 

Southern Indiana sandstone cliffs and wooded ravines, the 

August 20-22, 1970, The first stop was nei 
National Forest, an unusual deciduous 
characteristic of coniferous woods, such a 


American Fern Society 45 

tain Laurel, and Partridge Berry. There were abundant plants of 
Asplenium platyneiiron growing in open, grassy sites leading up to 
the woods. Pteridium aquilinum var. latiusculum, and Equtsetum 
hyemale also occurred in these open areas. One of the Gentians 
{Sahatia angularis), the False Foxglove {Aureolaria flava), and 
many composites were in good bloom. In the cool ravines were 
many plants of Botrychium virginianum and B. disseclum f, 
ohliquum, Adiantum pedatum^ Diplazium pycnocarpoUy D. acrosti- 
choides, Alhyrnim Filtx-femina var. asplemoideSy and Thelypteris 
hexagonoptera. Several orchids, the showy Orchis spectabilis^ and 
Rattlesnake Plantain (Goodyera repens), as well as Bloodroot, 
Hepatica, Banebcrry, Anemone, and Trilliunis were evidence of 
the rich Spring flora in these woods. On the higher slopes of the 
ravines, among rocks fragmented from the cliffs, there were 
numerous plants of Polystichum acrostichoides, Dryopteris margi- 
naliSj and fewer of Z). spinulosa var. intermedia and Cystopteris 
fragilis. On the spectacular, sheer cliffs, ledges were covered with 
Polypodium virginianum, and a few leaves of P. polypodioides were 
found. The pocked faces of the sandstone, especially where mosses 
were established, provided pockets for Asplenium pinnatifidum 
and A, trichornaues. One of the rare Saxifrages, Sullivantia Sulli- 
va7itii, kept company with these Aspleniums and Camptosorus 
rhizophyllus. Some of the more energetic climbers who scaled the 
cliffs found, on the upper slopes, one of the uncommon ferns of 
Indiana, Dennslaedtia pnnctilobida. 

. On the afternoon trip to Pioneer ^Mother's Memorial Forest, 
near Paoli, many ferns of the ravine w^ere seen again. This woods is 
one of the few remaining stands of primitive hardwoods in the 
state, with fine specimen trees of White Oak, Walnut, and Tulip 
Popular. There were several unusual orchids here— Lily Tway- 
blade {Liparis JiliiJoUa) , Puttyroot {Apledrum hyemale), and 
flowering specimens of the Nodding Pogonia (Triphora triantho- 
ph ova) . 

During the evening program, at our motel, in Jasper, RoUa 
Tryon reviewed the recent glacial history and vegetation of the 
region, and discussed the species that were collected during the 

American Fern Joubnal 

Volume 71, Plate 9 


^ -■ 







f -^^ 



.■ -( 





-^ > 


.. i". . _ _ _^ 



^ :;#. 

-^ ' 


Standing left to eight: Neill Hall, James .Montgonery, Jeanette 
Oliver, William and Garnet Meindbrs, Benton Stidd, Gregory AxNDER- 
SON, Donald Huttleston, John Clements, Susan and Gerald Gastony, 
Robert Lommasson, ^Iildred Faust, Rolla Tryon, James Maysilles, 
Mrs. R. Benedict. Victor Ribs, Lawrence Dillon, W. E. and Mrs. 
BUKER, Eva Sobol; Seated: Barbara Job Hoshizaki, Jeanette Gross, 
^LiNOR Henry, Lynda Cole, LeRoy Henry, Alice Tryon, Amy Mason, 
Betty Dillon, Ralph Benedict. Photo taken by Steve Wunderle. 

American Fern Society 47 

day. Neill Hall reported on the activities of the spore exchange 

and requested spores of any species, to replenish older supplies. 

Donald Huttleston spoke of the living fern collections at Long- 

■ wood Gardens, which is actively acquiring living collections from 


all parts of the world. Mrs, LeRoy Henry showed her colored 
movies and commented on the 19G7 Fern Foray, in Texas, and the 
1969 International Botanical Congress, at Seattle, including some 
of the attractive gardens of the area. The film had many good 
views of Fern Society members as well as habitats in Texas ana 

The second day's trip was to -NlacB ride's Bluff, another sand- 
stone cliff area, near Shoals. Woodsia ohlusa was abundant on the 
open road banks growing with the blue-flowered Lobelia siphilitica. 
Great colonies of Polypodkim polypodioides and Camptosorus 
rhtzophyllus covered boulders below the cliffs. A few plants of 
Cysfopteris hulbifera were found on the cliffs and one of the un- 
common ferns of Indiana, Cheilanthes lanosa, occurred on pre- 
carious ledges. Plants of Asplemum pinnatijidum and A, tricho- 
manes grew in association in pockets on the cliff faces. The rare 
hybrid between them, a fern new to the state, was a most appro- 
priate find by Gerald Gastony, a new pteridologist on the staff 
at Indiana University. 

An extra stop was made at the Martin State Forest, near Shoals, 
for several of the group who wanted to see Dryopteris goldiana. 

We were especially pleased to have among the thirty-three 
members and guests attending the Foray, several Hoosiers, 
enthusiastic and knowledgeable about their flora including, 
Gregory Anderson, John Clements, Lynda Cole, Amy and James 
2^Iason, Jeanette Oliver, and especially James Maysilles, of Han- 
over College, who arranged for our comfortable lodging and a 
fine picnic lunch. We are much appreciative of his help and that 
of Rolla Tryon in guiding us to some of their favorite haunts in 
southern Indiana.— Alice F. Tryo?;, Gray Herbarium, Harvard 
University, Cambridge^ Mass. 02138. 

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Office, as established under the act of October 23, 1962, Section 4369, Title 
39, United States Code, the following statements are published. 

Title: American Fern Journal 
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September 30, and December 31) 
Location of Office of Publication (Printers): 3110 Elm Avenue, 

Baltimore, Maryland 21211 
Location of Business Office of Publishers (Not Printers): Dr. LeRoy 

K. Henry, Section of Plants, Carnegie Museum, Pittsburg, 
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Publisher: American Fern Society, Inc., Section of Plants, Carnegie 

Museum, Pittsburg, Pa. 15213 

Editor: Dr. David B. Lellinger, Department of Botany, Smith- 
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Managing Editor: None. 

Owner: American Fern Society, Inc., Section of Plants, Carnegie 

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RipoB, WiseoBslii S4971 

Vol. G1 April-June, 1971 No. 2 

American Jfern Journal 


PtiblUhed hr th* 









Lindsaea (Schizoloma) ensifolia Swartz in Hawaii. W. H. Waoneb, Jr. 49 

The Fern Collections in Some European Herbaria, VU 

C. V. Morton 59 

An Appendageless Psilotuin. Introduction to Aerial Shoot Morphology 

Al-BERT S. RorFFA 75 

Vascularization of Fern Leaves 

Robert C. Lommasson and C. H. Yousg, Jr. 87 

Shorter Note : A Wrongly Localized Species of Pyrrosia. 94 

Recent Fern Literature 


Missouri BoTANICAi: 

JUL 16 1971 

Garden L-ibrary 

®j)e American jFern ^otitty 

Council for 1971 

Wabren n. Wagner, Jr., Department of Botany, University of Miohigan, 
Ann Arbor, Michigan 48104. President 

John T. Mickel, New York Botanical Garden, Bronx Park, Bronx, New 
York. 10458. Vice-President 

Richard L. Hauke, Department of Botany, University of Rhode Island, 

Kingston, Rhode Island 02881 Secretiry 

LeRot K. Hinht, Division of Plants, Carnegie Museum, Pittsburgh, Penn- 
sylvania 15213. Treasurer 

David B. Lbllinoer, Smithsonian Institution, Washington, D. C. 20560. 

Editor -in-ChieJ 

iiationat ^ocietp J^epceasentatibes 

Warren H. Wagner, Jr., University of :Michigan A.A.A.8. RepreaerUative 
Rolla M. Thyon, Jr., Harvard University A.I.B.S. Repreaentalive 

lamcrican iTcrn STournal 


David B, Lellinger Smithsonian Institution, Washington, D. C. 20560. 

G. \ . Morton Smithsonian Institution, Washington, D. C. 20.')C0. 

Rolla M. Trton, Jr. 

^ ^ ^ Gray Herbarium Harvard University, Cambridge, Mass. 02133. 

Ira L. Wiggins.... Dudley Herbarium, Stanford University, Stanford, Calif. 

An illustrated quarterly devoted to the general study of ferns, owned by 

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Changes of address, applications for membership, subscriptions, orders 
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Subscriptions $5.50 gross, $5.00 net (agency fee $0.50) ; sent free to mem- 
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1971, issue of the American Fern Journal was indicated at the top 
of the front cover, at the top of page 1, and at the right hand top 
corner of the plates, as being "Vol. 71." The correct volume num- 
ber is 61. Please correct your copy in order to avoid problems of 
bibliographic citation in the future. 

^mert can Jfern Journal 

Vol. 61 April-Juxe, 1971 No. 2 

Lindsaea (Schizoloma) ensifolia Swartz in Hawaii 

W. H. Wagner, Jr.^ 

Tlic fenis and other pteridophj^tes of the Hawaiian archipelago 
inchide approximately 180 species, of which 20 were naturalized 
since isr)0, liaving been introduced by man (Wagner, 194S, 1963). 
The islands have received an enormous amount of exploration by 
botanists over the years, but discoveries are still being made. 
Today we tend to assume that all of the major indigenous genera 
and species groups have already been found. Xevertheless, I wish 
to report a wholly distinctive species of fern not heretofore col- 
lected in the islands. As will be described below, the plant in 
(luestion grows in a rather unUkely habitat in association with other 
species which more or less resemble it.^ The fern could be readily 
overlooked; in fact, it was almost an accident that we noticed 

it at all. 

On December 16, 1969, a group of botanists and ecologists 
visited the Puhimau ''Hot Spot,'' an area along tlie Chain-of- 
Craters Koad in Hawaii Volcanoes National Park, just south of 

^ Research supported by NSF Project ' 
tern of Fern?5. 1 wish to acknowled^^e th 
Gressitt, K. U. Kramer, D. B. Lellinger 

2 One such species is Pteris vittata L. AUhough adult plants of this species 
are obviously different from Lindsaea ensifolia, the juveniles are more difficult 
to distinguish. They differ in having all of their pinnae short to even orbicular 
in the youngest stages and in being pinnate at early stages. Those of U ensijo ta 
are ternate and perhaps even snnple at early stages, and become pinnate 
only at later stages. — D.B.L. 

Volume 61, No. 1, of the JorRXAL, pp. 1-48, was issued March 23, 1971. 

American Fern Journal 

Volume 61, Plate 10 

-^^4i6i-i^-\ ■■^;=r':=-^'*^ 


appearance of vegetation and plumes of hot steam. Fig. 2, Obseuvkb 




Puhimau Crater (briefly described by Doty and Muoller-Doniboi.s, 
1966, pp. 322-323). Surface and steam vent temperatures here 
run from 4S°-89° C, and the ground is warm to the touch. The 
roots of many of the pUmts, inchiding the conspicuous grasa 
Andropogon virginicus and scattered small trees and sln-ubs of 
Metrosideros are only at the surface and do not penetrate into the 
hot earth. For this rea-son the plants are not anchored and can 
be literally moved around and re-arranged like pieces of furniture 
on the white, ash-like soil. Plumes of steam arise in an eerie 
fashion in this other-worldly habitat (Fig. 1,2). Around the steam 
vents are dense growths of mosses and ferns {Plate 11). The ferns 
are mainly Swordferns, Xephwlepis exaltata, but occasional 
patches of phioglossiun nudicmde occur among the mosses. The 
Swordferns tend to become progressively smaller the closer they 
grow to the steam, as shown in the lower half of Plate 11. 

While we were examining the Swordferns around the steam 
vents we first noticed small plants that seemed especially in 
texture to be different from the juvenile plants of other fern species 
known to occur in the Kilauea area. Careful examination of these 
plants and their associates finally revealed older stages of this 
fern, which we realized was a species unreported in the islands. 
Colonies of it, usually vastly outnumbered by the associated 
and grossly similar-appearing Xephrolepis, were later located 
around several of the steam vents. Subsequently the plant was 
identified as Lwdsaea {Sch izoloma) eusifdia, about 2G00 miles 
from its nearest other localities in Samoa. 

lu the following jiaragraphs comments will be made upon the 
leaf structure, juvenile and mature, and the significance of the 

occurrence of this fern in Hawaii. 

The early heteroblastic stages {Plate 12) are different in appear- 
ance and structure from the mature stages in Plate 13. The earlier 
stages look similar to the corresponding juvenile stages of the Tree- 
fern Cihotium. But the little plants of Limlsaea are glabrous and 
much thicker-textured than those of Cihotium. The young stages 
have jagged, more or less lobed piima margins with up to two 
pairs of pinnae. The veins are free, except the larger specimens 

American Fern JouRNAii 

Volume 61, Plate 11 


Edge of Fumerole with Progressively More Depauperate Forms ok 
Dominant Nephrolepis exaltata (Photograph by Mueller-Dombois), 


(on the right side on Plate 12) which have more or less casual anasto- 
moses. These fronds are transitional to the mature forms shown 
in Plate 13. Holttum (1954, p. 342) in his discussion of this species 
and its relatives comments that: 

The main evolutionary trend in Schizoloma [a taxon 
that he upholds as a genus distinct from Lindsaea] is 
towards a simply pinnate (or even simple) frond with 
reticulate veins and a long fusion sorus. On this view, 
the common Malayan S. ensifolia represents an ad- 
vanced type . . . 

It is interesting to note, therefore, that the heteroblastic series of 
Lindsaea ensifolia leaves from HaAvaii seem to follow approximately 
Holttum's concept of the evolutionary trend in frond evolution, 
a possible case of recapitulation. 

The largest fronds of L. ensifolia that we were able to find are 
consistently trifoliolate (Plate 13), Their veins are copiously netted 
(for details of pattern see Wagner, 1952, fig, 24,6). The leaf 
margins of the mature fronds are nearly entire. We were able to 
find only a few fertile fronds, and these were only incompletely 
fertile, with the sori interrupted and widely separated. Efforts 
should be made to obtain more fertile fronds in order to determine 
whether the Hawaiian form has consistently discrete sori. 

Mature fronds were sent to Dr. K. U. Kramer of the State 
University of Utrecht, Netherlands, who has made extensive 
studies of lindsaeoid ferns in various parts of the world. In com- 
menting on these specimens (letter, March 4, 1970), he wrote: 

It is indeed strange that only trifoliolate leaves of L. 
msijolia have been found at the Hawaiian fumeroles. 
Offhand, without knowing the exact local situation, I 
tihould say that apparently the plants fail to reach a 
more mature stage with midti jugate lamina. In what 
I take to have been poor habitats in the mountains of 
e.g. New Guinea even simple forms have been col- 
lected, la the genus Lindsaea it is not uncommon to 
find fertile leaves even in small, evidently far from 
full-grown leaves. 

American Ferx Journal 

Volume 61, Plate 12 

i^wW^-HH^ ILVfLlxi ■-- '^ 

--^v::o:j -^^^v-i-^^^^-roi^Sv:- :-":V= >■=:-:■>■■-■ 






'"^'^H-^.-. .v;v;Vb';^-^ 



^ ■ ■ ■ ■- ' ^^ -_'_K'.-^iita-j'.-_- 

Heteroblastic Leaf Stages of Linds^aea ensifolia from Steam Vent 
Edges with Earlier Stages at Left. 


In another (■ommuiiication (January 7, 1970), Dr. Kramer stated 

The most intriguinjg; thing is perhaps that this is 
clearly ssp. ensifolia which has a frestern distribution 
(East to Micronesia and the Solomon Islands), not 
ssp. again which I have seen from Micronesia, Mela- 
nesia, and Polynesia (Fiji, Samoa, Tonga), which is 
the eastern subspecies. [For details of the contrasts, 
see Kramer, 1967a, p. 679]. 

As Lindsaea ensifolia was not encountered in Hawaii until 
December, 1969, one might argue that it had been introduced 
by man, either intentionally or unintentionally. I am, however, 
hesitant to accept this argument. This is not a plant likely to be 
carried by man. Unlike various naturalized ferns of Hawaii (cf. 
Wagner, 1948), L. ensifolia is a relatively unattractive fern and 
is not grown at all in horticulture, to my knowdedge. The area 
where it occurs in the fumeroles has been studied in some detail 
by botanists, it is true, but the appearance of the plant is such 
that it can be casually confused with imperfectly developed stages 
of other ferns, e.g., Cihotium or Nephrolepis. It can be overlooked 
readily in a tangle of dwarfed Nephrolepis fronds (Plate 11). The 
geographical separation of the Hawaiian plants from other Poly- 
nesian localities is very large (the nearest are Samoa, Fiji, and 
Rotuma, according to St. John, 1954), but such large disjunctions 
across the open seas are not uncommon among Pacific ferns. In 
fact, the geographical distribution is similar to the two other 
lindsaeoid ferns known in Hawaii, Lindsaea repens and Sphenomeris 
chineiisis. Fern spores are capable of long-distance dispersal 
because of their extremely small size. Current research suggests 
that a single spore has the potentiality of initiating a new colony 
of ferns by gametophytic self-fertilization. Fern gametophytes 
tend to be self-compatible according to the recent studies of 

Klekowski and Baker (1966). 


peculiar. However, L. ensifol 

is very commonly a sun fern, growing in open, lateritic savannas, 

American Fern Journal 

Volume 61, Plate 13 


- >>^^:«o '- 

-■^0 ^>^^>^:-:^-■-!^''><. ■:v:o:^^^■;■^>^^:■''^■>'----^^-'--> -:-- t^y-yj^yj^.-^ /^>-' 

r TT y — n n - — x- - -t— x — — - xtx - »i^ -xT — - rx— n-- 


Mature Fronds of Lindsaea ensifolia ftiom the Steam Vents, Care- 
fully Spread out in Pressing (left) and Pressed Directly Without 
Spreading (right). 


and capable of withstanding the heat of direct sunlight. Holttum 
(1954, p. 243) states that it "has tlie power to live in poor soil, 
in exposed as well as shaded places." In the Kilauea fumeroles it is 
associated with Ophioylossian midicmih, a common associate also 
in the savannas of ^Ticronesia (e.g., Guam) and Melanesia (e.g., 

Admiralty Islands) . 

The very brief list of lindsaeoid ferns of Hawaii has now been 
expanded from two to three. These are morphologically different 
from one another, but they do share rather similar, broad ranges. 
Limhaea repens (Bory) Thwaites belongs to subg. and sect. 
Odontoloma according to Kramer, and is interpreted by him 
(19fi7b, pp. 508, 5G9) as being "very variable, and a number of 
more or less limited infraspecific categories can be distinguished." 
He interprets the Hawaiian plant as var. macraeana (Hook. & 
Arnott) Kramer. Liudsaea ensifoUa belongs to subg. Limhaea sect. 
Schizohma according to Kramer (19t)7a). Sphenomeris chiitensis 
(L.) :\faxon was previously designated by most recent writers as 
S. chusana (L.) Copel., but the latter name is evidently incorrect 
(Kramer, 1967a; Fosberg, 19G9). All three of the Hawaiian linds- 
aeoid ferns range as far west as tropical Asia. 

Further field studies of the fumeroles of the Kilauea area are 
recommended to determine whether other still unreported fern 
species have been overlooked in these seemingly unlikely fern 
habitats. Outside of Hawaii, I am aware of only one previous re- 
port of a strongly disjunct fern population around hot effluents. 
T. AI. C. Taylor (1963) reported an unusual occurrence of the 
southern maidenhair fern, Adiantum capillus-veneris L., as follows: 

In British Columbia it is known from the runnels of the 
hot springs at Fairmont, north of Columbia Lake. 
Despite the fact that the nearest known colony is some 
hundreds of miles to the south, there is very strong 
circumstantial evidence to support the view that it is m- 
digenous at Fairmont and not an escape from culti- 

In the Hawaiian fumeroles there is no reason to believe that 
the disjunct ferns reported here are not also indigenous. Now that 

American Ferx Journal 


are found iii the steam vents of Kilauca and are apparently limited 
to them, interest in the investigation of these unusual habitats 
should be stimulated. 


DoTv, M. S. and D. Mueller-Domhois. 1966. Atlas for Bioecology Studies 

in Hawaii Volcanoes National Park. Hawaii Bot. Sci. Paper 2. 
iii + 507 pp. 

Fosbeug, F. R. 1969. The nomenclature of Sphenomeri.s chinensis [Filic.]. 

Taxon 18: 596-598. 

HoLTTi M, R. E. 1954. Ferns of Malaya. Gov't. Printing Office, Singapore. 

iv + 643 pp. 

Klekowski, E. J., Jr. and H. G. Baker. 1966. Evolutionary significance of 

polyploidy in the Pteridophyta. Science 153: 305-307. 

Kramer, K. U, 1967a. The lindsaeoid ferns of the Old World. I. Xew Cale- 
donia. Acta Bot. Neerl. 15: 562-584. 

;• 1967b. The lindsaeoid ferns of the Old World. II-IIL Notes on 

Lindsaea and Sphenomeris in the Flora Maleisiana Area. Blumea 

St. John, H. 1954. Ferns of Rotuma Island, a descriptive manual. Occ. 

Papers Bishop Mus. 21 : 161-208. 
Taylor, T. M. C. 1963. The Ferns and Fern-allies of British Cohnnbia. 

British Columbia Prov, Mus. Dept. Recreation and Conservation 

Handbook No. 12. 172 pp. 

Wagner, W. H., Jr. 1948. Ferns naturalized in Hawaii. Occ. Papers Bishop- 

Mus. 20: 95-121. 


. 1963. Pteridology in Hawaii. Hawaiian Bot. Soc. Newsletter 2: 


Matthaei Botanical Gardens, University of :Michigan, 
Ann Arbor, Michigan ISlOo. 

Ferns in European Herbaria 59 

The Fern Collections in Some European Herbaria, VH 



There were doubtless some herb gardens iu Brussels from very 
early times, but the first formal association of gardeners was that 
of the Confrerie de Sainte-Dorothee formed in 1G50 and named in 
honor of Saint Dorothy, supposed for some reason to be the patron 
saint of flowers. This venerable institution persisted until the 
French occupation of Belgium in 1794. In 1822, a number of the 
former members reconstituted the society under the more appro- 
priate name of the Societe de Flore, and it still exists today as a 
part of the Societe Royale Linneenne et de Flore. 

Under the Frencli occupation the government established a 
botanical garden in Brussels in 1797, which was located in the 
grounds of the palace known as the "Ancienne Cour." This garden 
was intended primarily for the instruction of students in the 
Ecole Centrale. It had a considerable number of rare plants 
arranged by the Linnacan system. 

In 1825, four botanists, headed by J. B. Meeus, approached the 
burgomaster for permission to found a company to build a new 
botanical garden. An area of about 15 acres in a good part of 
Brussels on the newly proposed Rue Royale was at that time 
the site of a market-vegetable garden. This was bought by Meeus 
for 43,537 francs. The society was formally approved by a royal 
decree May 28, 1826, under the name Societe Royale d'Horti- 
culture des Pays-Bas. A competition was held for the design of a 
building, and that by the French painter Geneste was chosen. 
The building, still standing today, is a fine example of classical 



^^L ^1 ^L ^B ^L I ^m ^F H ^L ■ ^1 ^1 H H ^H B ^1 ^1 ^H H H B B A ^k ^^L ^f ^h ^H ^B ^^k ^v_p^^^p ^H ^^L ^^L ^^^^^^h '^^^r ^T ■ ^m ^^p 

houses were already finished in 1827, ready to receive the plants 
from the garden of the Ancienne Cour, which was being abandoned 

00 American Fekx Joukxal 

after a disastrous fire in 1820 and the conseciucnt reconstruction 
of the court. 

Ill 1814, near the end of the Napoleonic period, the powers 
formed the United Netherhuids under ^Yilliam T, but this union 
of the Dutch and Belgians was an uneasy one due to economic 
problems and the divisive influences of religion and language. The 
Belgians broke away in 1830 and Dutch troops sent in to quell the 
uprising lodged in the new greenhouses, causing great damage. 
There perhaps was no herbarium in the botanical garden, the only 
herbarium being the Rijksherbarium, at that time in Brussels 
under the directorship of Blume. When the Dutch withdrew in 
1830 they took along the state herbarivmi to Leiden, where the 
Rijksherbarium still is, a circumstance that shows how much 
herbaria may be valued even in times of war and stress. Later, the 
new king of the Belgians, Leopold I, of Saxc-Coburg, gave the 
garden the collections of herbarium specimens that belonged to 
the royal court. After the independence of Belgium, the name was 
officially changed to Societe Royale d'Horticulture de Belgique. 

The new garden was financed by the selling of shares, for which 
the stockholders were to receive dividends. The city and govern- 
ment agreed to a subsidy, but the amount was small, and so the 
only way that the society could make money was by the sale of 
plants. Thus almost by necessity the garden was forced to have 
plants that would grow easily in Brussels and which would sell 
well, and this consequently led to the neglect of strictly botanical 
work. Even so, severe financial troubles in 1841 caused a move to 
dissolve the society, which was in debt for over 203,417 francs. 
The salvation of the botanical garden was mostly the work of one 
of the giants of Belgian botany, Barthelmy C harles Joseph 
Du Afortier. He was born in Tournai April 3, 1797. He did not 
receive any formal training in botany, but nevertheless devoted 
himself to It wholeheartedly from about the age of 19, when he 
began collecting plants in 1810. He married also at the age of 19 
and was devoted to his vnfe, by whom he had eight children, foi 
the next 60 years. He must have had independent means and some 
social status because he worked independently studying and 

Ferns ix European Herbaria (51 

writing on botany until 1831, at which time he was elected to the 
Chambre dcs Represcntants, the equivalent of our House of 
Representatives. He continued to be reelected for the next 47 
years until the time of liis death in 1S78 at the age of 8L 

In the early nineteenth century the Linnaean system was still 
dominant but that of Jussieu was gaining adherents. Du ^lorticr 
considered the Linnaean system analytical and that of Jussieu 
synthetic, and he wished to amalgamate them, an idea that seems 
rather impossible on the face of it. However, as Crepin remarked, 
because of his lack of training he did not hesitate to discuss the 
most difficult problems in botany and zoology. When he was 24, 
Du IMortier personally published his "Commentationes Botanicae" 
(1822), in which some of his ideas were expressed, a book that is 
said to have caused a sensation in Belgian botanical circles. The 
part that is remembered now is the h\st chapter, in which the old 
cumbersome genus of hepatics Jun.,ermannia was broken up into 
several, a view that is certainly now considered correct. Du 
Alortier's work was original, and he was doubtless much dis- 
turbed to find out in 1831 that Raddi had anticipated him by 
publishing a similar segregation of Jun^ermannia in an obscure 
Italian publication in 1820. Nevertheless, he continued working 
on hepatics, publishing his "Sylloge Jungermannidearum Europae" 
in 1831 and more than 40 years later the complete monograph 
"Hepaticae Jungermannidearum Europae" (1871). 

Du Alortier worked also on phanerogams, especially Scrophula- 
riaceae, Batrachium, Salix, and Gramineae. His "Florula Belgica 
Prodromus" (1827) was intended as the forerunner of a complete 
flora of Belgium, something not in existence then, but he never 
finished this. Before writing this, Du Mortier visited the Linnaean 
Herbarium, then in London in the custody of Sir James Edward 
Smith, to study types, one of the ratlier few early botanists who 
availed themselves of this privilege. At the same time he also made 
an expedition to Scotland to see the elder Hooker. [In 1826, a trip 
to Scotland probably really deserved the term "expedition."] 

At the International Botanical Congress in Paris in 1867, 
Du Alortier presided at the sessions on nomenclature, the 

()2 Aaierican Fern Journal 

principal interest of the Congress and one tliat took up over half 
of its time. In 1872 he was made a count by Leopold II. The 
medal struck for him has below the portrait the words ^'Felix qui 
potuit rerum cognoscere causas'' [Happy is he who can know the 
(pauses of things]. On his 81st birthday in 1878, his statue was 
unveiled in the botanical garden, a bust that is still there in a 
place of honor. 

Du Mortier, as commissioner in charge of scientific establish- 
ments, was able to persuade the government to double its sub- 
vention of the botanical garden, provided that the society would 
renounce its right to dissolution without the consent of the govern- 
ment. Ihe society was authorized to sell some of its grounds oi) 
the side of the present Gare du Nord in order to pay its debts. A 
period of prosperity then began, especially after H. G. Galeotti 
was selected as Director in 1853. 

Henri Guillaume Galeotti was born September 10, 1814, in 
Paris, Frame, of Italian parents. He was educated as a geolo- 
gist, and published something on the geology of the region of 
Brabant at the early age of 20. Tliis brought him to the attention 
of the Vandermaelen Brothers, the proprietors of a geographical 
institute and museum in Brussels. They sent him in September, 
1835, to ]Mexico to study the geology and to collect plants, both 
living and dried, which Galeotti did vigorously, assembling more 
than 8,000 immbers of herbarium specimens in five years. He 
began his collecting in the rich region of Alirador and Jalapa in 
Veracruz. He was perhaps the first botanist to climb the high 
mountains Orizaba and the Cofre del Perote. In July, 1830, he 
collected in Real del IMonte, Hidalgo, with the German botanist 
C. Ehrenberg. The next three years were spent on the plateau of 
Mexico and in the west. In 1839 he went to the southern state of 
Oaxaca, -where he obtained his richest collections. Before returning 
to Europe in 1840, he established a botanical station in Mirador 
which was used by later collectors. 

On his return to Belgium he was made Administrator of the 
Societe Royale de Flore and later editor of the Journal d'Horti- 
culture Pratique. He lived in Louvain, where he became a natu- 

Ferns ix European Herbaria 63 

ralized Belgian in 1S48. His wonderful collections were studied by 
some of the foremost botanists of the time, the cacti by Lemaire, 
the grasses by Trinius, and the orchids by Achille Richard. Many 
of the ferns and phanerogams were published on by the Professor 
of Botany in Louvain, Martin Martens (1797-1SC3). Galeotti 
himself although sometimes a co-author seems not really to have 

been himself a taxonomist. 

Unlike most early collectors, Galeotti collected many duplicates 
and sold them widely, and so they are in most of the large herbaria. 
Dr. R. Tournay, documentation chief in Brussels, tells me that the 
Galeotti specimens belonging to :\rartens were held by the Martens 
family until 1932, when Pierre Martens, greatgrandson of ^1. 
Martens, gave them to Brussels. However, the "Index of Collec- 
tors" of the "Index Herbariorum" (Regn. Veg. 9, 11(2): 214. 1957) 
indicates that the original and largest set (7,000 specimens) isstill 
in Louvain, which is a matter that ought to be looked into. Galeotti 
collected only in Mexico and a few specimens in Cuba but is often 
cited as the collector of plants from Brazil, Venezuela, and Colom- 
bia, but these plants were actually collected by Linden, and Gale- 
otti was merely the distributor of the Linden plants. In 1853, he was 
appointed Director of the Jardin Botanique de Eruxelles, a post 
that he retained until his death from tuberculosis in 1858 at the 
age of 44, a relatively early age considering that as a group Belgian 
botanists seem to be exceptionally long lived. During his adminis- 
tration an herbarium was started with Galeotti's ow^n collections 
as a basis, to which were added those of Linden, Funck, SchHm, 
Ghiesbreght, Lindig, and the complete Brazilian herbarium of 
Peter ( laussen. The botanical library was also enlarged. 

After the death of Galeotti, A. J. Schram became the Acting 
Director, and J. E. Bommer w^as appointed as conservateur (cura- 
tor). Mr. Schram may not have had the best judgment. The erec- 
tion of an aquarium was a costly mistake. When Schram left, 
Bommer was apparently the Acting Director. The necessary re- 
placement of the heating equipment in 1862 caused some financial 
embarrassment, so that again the stockholders received no divi- 
dends. The administration tried without success to get additional 

<>-4 Ameuican Fern Joukxal 

subsidies from tlie city and government. In I8G0, the garden had 
tempting offers of up to 6,000,000 francs for permission to allow 
roads to run through the garden, and so there was again a deter- 
mined move to dissolve the society. Again Du ?iIortier, who was 
still powerful in the government, came to the rescue. He per- 
suaded the government to buy the garden for 1,000,000 francs, 
not a large sum even at that time for such a valuable property. 
However, it was doubtless large enough for the shareholders to 
receive their equity. The garden was then renamed the Jardin 
Botanique de I'Etat [in Flemish, the Rijksplantentuin], a title that 
it retained until 1967, when the name was changed for some 
unexplained reason to the Jardin Botanicpie National de Belgifjue 
[Nationale Plantentuiu van Belgie]. 

The official establishment of the Jardin Botanique de I'Etat in 
1870 was followed a year later by a most propitious occurrence. 
The famous Bavarian botanist Karl Friedrich Philipp von ISIartius 
(1794-1868) had assembled a very large private herbarium, in 
excess of 113,000 specimens, which was offered by the family to 
the government of Bavaria for purchase, but the government 
very short-sightedly refused, because it had already the Brazilian 
herbarium of Alartius and Spix. The family therefore wrote to 
Anton Spring, the monographer of Lycopodium and a former 
student of Martius, who formed a committee consisting of him- 
self, Alphonse De Candolle, and August Eichler. De C'andolle 
wrote a flattering letter to Du Mortier, ending: "J'ose insister, 
parce qu'il est bien rare qu'un President de Societe botanifiue et 
uu botaniste consomme soit en meme temps un politique 
d'un pays. Ce sent dcs conjonctions aussi rare (lue celles de certain 
corps celcstin. La Science doit en esperer beaucoup." Du Mortier 
was persuasive as usual and the Belgian government purchased 
the .Martius Herbarium in 1870 for 32,000 francs, a ridiculou.slv 
small sum even for those days for an herbarium containing thou- 
sands of types and unique specimens. The herbarium was delivered 
to tlie state botanical garden in 1871, where it forms the real basis 
today of the general herbarium. The collections in this herbarium, 
which were enumerated by Eichler in his paper on the Herbarium 

Ferns in European Herbaria 65 

Martii, are far too numerous to list here. They inchulc liistoric 
specimens from all parts of the world. 

The state garden has had very few Du-ectors since its establish- 
ment a century ago, principally Crcpin (1876-1901), Durand (1902- 
1912), De Wildeman (1912-1931), Walter Robyns (1931-1966) 
and Fernand Demaret (1966-present). After Provisional Directors 
Bommer and Dupont, Frangois Crepin was appointed Director in 
1876. Crepin was the antithesis of Du :\Iortier, not a politician and 
outgoing public figure but a retiring man content with his work 
with plants. He was born in Rochefort, October 30, 1830, and 
never received a degree or any formal training in botany. While he 
was working as a postman at the age of 18 he began botanizing in 
his spare time, and at 20 quit the post office to devote himself 
wholly to botany without any sponsor or any hope that he would 
ever have an official position. For the next ten years he collected 
and studied the plants of Belgium, and his resulting book, the 
"Manuel de la Flore de Belgique" published in 1860 immediately 
established his reputation. Because of its compact size and keys, 
the work went through several editions and enlargements and was 
the standard field-guide for two generations of Belgian botanists. 

His friendship with Louis Van Houtte stood him in good stead. 
Van Houtte had been employed in the :\linistry of Finance. 
However, when one day he was requested to undertake an im- 
portant mission over the week-end he replied that it was impossible 
since he had a date to see the flowering of a rare cactus. The reply 
did not please the Minister, and from then on Van Houtte was 
free to devote himself wholly to observing plants, something 
indeed fortunate for botany since he became one of the foremost 
protagonists of horticulture and botany in Belgium and indeed 

in all of Europe. 

In September of 1801, Michel Joseph Scheidweiler (1799-1861), 
then in the Institut Horticole of Gentbriigge, died leaving vacant 
the chair of botany and agronomy that Planchon had for a time 
occupied. Van Houtte was able to get Crepin this professorship 
which meant that Crepin immediately had to courses 
in botany, a task that he did creditably. However, he had little 

66 American Fern Journal 

interest in garden plants, and Van Houtte's continuous impor- 
tunities to name his greenhouse plants caused a break in their 
friendship. Therefore, Cr6pin left Ghent in 1871 to become the 
curator of the paleontological section of the INlusee Royal de 
THistoire Naturelle de Bruxelles. The government had purchased 
a collection of plant fossils from E. Coemans, and Crepin was 
delegated to name them. In 1872 he was elected to the Academie 
des Sciences de Belgique and in 1876 became the Director of the 
Jardin Botanique de TEtat. 

The staff near the beginning consisted of Bomnier, Cogniaux, and 
Marchal, to which were added sometime later Delogne, T. Durand, 
and De Wildemanj a distinguished staff indeed. Tlie working room 
was at the time the large room now containing some offices at the 
sides and a wood exhibit in the center. Crepin sat in the middle at 
a big table on a dais where he could survey the work of his staff. 
On his left at the most primitive kind of tables barricaded by 
stacks of books and cartons of herbarium specimens sat Bommer, 
Elie Alarchalj and the bryologist Charles Delognc, and on the 
right Sonnet, Vindevogelj Lubbers (the Head-Gardener), and T. 
Durand. De Wildeman, as only a volunteer at this time, sat in 
1SS7 at a table a little to the rear. Working conditions must have 
been difficult, for at first there was no illumination other than the 
natural light from the windows, and Brussels is not notably bright 
during the long Avinters. Only in later years was gaslight installed. 
Yet the amount of work accomplished was enormous. Crepin 
himself worked on the genus Rosa^ a task on which he spent 40 
years, and on which he published over 1,500 pages, but even so 
his work was never fully completed. His rose herbarium of over 
40,000 sheets is kept today as a special unit. 

Crepin retired because of poor health in 1901 and died In 1903. 
He was described on a field-trip in 1893 by a journalist as a vigorous 
sexagenarian, a confirmed bachelor, with a voice sweet and low, 
who looked like a retired army major except for his modest and 
reserved attitude, always with his eyes on the ground as if looking 
at something [which does not seem at all strange to a botanist]. A 
statue of him was erected in 1904 in his birthplace in Rochefort 

Ferns in European Herbaria 67 

in the square named in his honor, the Place Franyois Crepin. For 
a portrait of Crepin see the biography in Bull. Jard. Bot. de TEtat 
Brux. 1 : 281-317. 1905. Fascicle 2 of volume 9 of the same journal 
is devoted to the celebration of the centenary of Cr6pin^s birth. 

Two of the staff members deserve special mention. Cdlestin 
Alfred Cogniaux (1841-1916) was born in Robechies. He was 
appointed Aide-naturaliste in 1872 in the newly organized Jardin 
Botanique de TEtat, where he remained for a number of years. 
He was a prolific worker on three difficult plant families, writing 
the Cucurbitaceae (1878), Alelastomataceae (1883-1888), and 
Orchidaceae (2037 pp., 1893-1906) for Martius' "Flora Brasil- 
iensis/' and the Cucurbitaceae (602 species, of which 221 were 
new, 1881) and the ^Melastomataceae (2,730 species, of which 792 
were new, 1891) for DeCandolle's "Monographiae Phanerogamae/' 
Finally in his later years, 1909-10, he published an account of the 
orchids of the West Indies in Urban's ''Symbolae Antillanae." 
His valuable herbarium was sold to Brussels. 

Elie Marchal was born :\Iarch 1, 1839, at Wasigny, France, of 
Belgian parents. In 1871, he was chosen by Bommer as Aide- 
naturaliste in the Jardin Botanique de TEtat, where he worked 
on mosses and fungi. He was in charge of the plantings in the 
garden, where he established sections for ornamental plants, 
edible plants, medicinal plants, and plants of technological im- 
portance, all these in addition to the plants of primarily botanical 
interest. This was in opposition to some prominent Belgian botan- 
ists like Kickx, Morren, and Martens, who wanted only plants of 
botanical interest. He was ahead of his time in insisting that all the 
plants were properly ticketed with their names and acquisition 
data. In his later years he worked on Araliaceac, and prepared 
the account of this family for the ''Flora Brasiliensis.'' For reasons 
of poor health he retired in 1899, but nevertheless lived up until 
1923, when at the age of nearly 84 he was still interested and 
working on sexuality [of the mosses]. 

The next Director was Theophile Alexis Durand, one of 
Belgium's outstanding botanists, who was born in 1855 in St. 
Josse ten Noode. He explored the region of Liege and published 

OS American Ferx Journal 

his first paper on the flora in 1874 at the age of 19. At the sug- 
gestion of Elie Marchal he went to Switzerland in 1877 to study 
the alpine plants, and published a series of papers (1881-1887) with 
his friend Henri Pittier, at that time Professor of Botany at 
Chateau d'Oex. In 1879 he started work as a volunteer in the 
Jardin Botanique de I'Etat. By his continuous work and the care 
he gave to all the tasks confided to him he won the regard of 
Crepin and was appointed to the staff as Aide-naturaliste in 1881, 
and later became the curator. At first he worked on the bryophytes 
of Belgium with Delogne, but soon became interested in com- 
piling an index of genera, the result being the "Index Generum 
Phanerogamorum," a work that made him famous and one 
that is still most useful to the herbarium botanist. He then col- 
laborated with B. D. Jackson in preparing the first supplement to 
the "Index Kewensis." In the meantime he continued fioristic 
work with Pittier on the "Primitiae Florae Costaricensis." In 
1895, at the request of the government, it was decided that the 
garden should undertake the study of the flora of the Belgian 
Congo, and Crepin delegated Durand and De Wildeman to the 
task. This collaboration resulted in the formation of the "Annales 
de Uu&6e du Congo" and the publication of many important 
papers on the plants of that region until eye trouble forced his 
retirement in 1908. He died in 1912.i 

The next Director, one of the brightest lights of Belgian botany, 
was Emile De Wildeman, who was born in Brussels, October 19, 
1866, where he died July 24, 1947. He became a volunteer worker 
in the botanical garden in 1887 and was taken officially on the 
staff in 1890, where he remained for 41 more years of service 
before retiring in 1931. At the time he was one of the first students 
of the famous professor Leo Errera (the formulator of what is 
called "Errera's Law" that a cell membrane tends to assume the 
same form as a weightless fluid layer would assume under the same 
conditions) , who had just started a botany department in the 

4r2v'i-vv^i-''^i'Qr]'^i^- ^ Wildeman, see Bull. Jard. Bot. de I'Etat Brux. 
AmPnV«n Fi'i« ir ^^'^ Durand is not to be confused, of course, with the 
S^esTnow in ^riF '"''''' ' ^^ ^^^ ^'*°"*' herbarium containing Rafinesque 

Ferns in European Herbaria 69 

University of Brussels in 1891. Do Wildcman received bis doctorate 
in 1892 with a thesis on cell division. In liis early years lie woiked 
and published on algae, but in 1895 Crcpin decided to charge him 
with the preparation of a flora of the Belgian Congo, a task on 
which he spent the rest of his life. The herbarium collections that 
had been accumulated by the Etat Indepcndant du Congo were 
turned over to him, the begiiuiing of the present tremendous 
collection of Congo plants numbering more than 500,000 sheets 
in 194S. De Wildeman became the foremost authority on the 
flora of the Congo, and was very likely the last botanist to describe 
thousands of new species [because there are no more completely 
unknown parts of the globe]. It was perhaps not necessary to 
describe quite so many, because recent studies have shown that 
De Wildeman did not always take into account the species that 
had been described from other parts of tropical Africa, and also 
had perhaps too narrow a species concept, but even so most of 
his species are good ones. He was one of the most prolific workers 
ever, with a bibliography of more than 1,400 titles, ranging from 
brief notes to large books. His final work, entitled rather sadly, 
"Sterilite ou Viellissement et Disparition des Especes Vegetales" 
was published in two volumes a year after his death at the age of 
80 in 1947. He was the Secretary-General of the International 
Botanical Congress that was held in Brussels in 1910. In 1912 he 
became Director of the Jardin Botaniquc de I'Etat." Young 
botanists and perhaps especially the old too) should ponder his 
words: "Ne cherchons pas a faire oeuvre definitive! Ne nous ber- 
gons pas du fallaceiux espoir de produire des travaux sans de- 
fauts. La perfection n'est pas de notre monde!" 

During the regime of W. Robyns, the next Director, it became 
apparent that the old building and greenhouse were inadequate 
and that the garden was suffering from the city air. It was there- 
fore arranged in 1938 that the garden be moved to the Domaine 
de Bouchout, in the country near :\Ieise, a few kilometers north of 
Brussels. This is a large estate of over 200 acres, with a lake and 
an imposing chateau, formerly a royal residence. The advent of 

' For a biography, see Bull. Jard. Bot. de I'Etat 19: 1-31. 1948. 

70 American Fern Journal 

the war in 1939 naturally stopped any move to the new quarters. 
About 10 years later work was started, and now the huge green- 
houses are finished and all the plants moved from the old ones. A 
magnificient new administration and herbarium building is 
awaiting occupancy. The new garden is not so accessible as the 
old one in the center of Brussels, but is well worth a visit. In 
addition to the greenhouses the grounds themselves contain many 
fine old trees dating back a century or more. Certain areas are 
being left in a natural state for the wild vegetation. The new garden 
may be reached rather easily by taxi or bus. There are however, I 
believe, no accomodations liearby where visitors can stay for 
longer periods. 

Robyns continued with De Wildeman's w^ork on the Congo, 
publishing volumes on the grasses and a 'Tlore des Spermato- 
phytes du Pare National Albert'' in three volumes (1948-1955). 
He also started the publication of the ''Flore de Congo Beige et du 
Ruanda-Urundi," of which ten volumes appeared between 1948 
and 1963. This work is being continued, but now in the form of 
separate fascicles for the different families, which can thus be 
issued as they are ready without any set taxonomic sequence. 

The present Director, Fernand Demaret, an authority on the 
bryophytes of the Congo, first came to the garden in 1936 as an 
assistant and became director in 1966. 

September 1970 being the centenary of the establishment of the 
garden as a government unit, the garden held a formal celebration 
of the event. Botanical institutions throughout the world were 
invited to send representatives for the ceremonies. Since I had 
been in Brussels in June of that year, I did not feel that I could 
attend, but I understand that the event was highly successful in 
every way. 

Ferns in Brussels 

The principal pteridologist in the early years was Jean Edouard 
Bommer, who was born November 17, 1829, in Brussels. He had 
no formal education in botany, but w^as a hard and dedicated 
worker, at first employed in the Etablissement Vandermaelen, 

Ferxs in European Herbaria 71 

but by 1856 he was well enough known that he joined the staff of 
the botanical garden in Brussels as Aide-naturaliste, and after 
Galeotti^s death he was the curator and later Provisional Director. 
Although Bommer worked on various groups of plants, he was 
mostly interested in ferns. His first paper was on the hairs of ferns, 
a subject that has been somewhat neglected but now realized to 
be of fundamental importance (Bull. Soc. Roy. Bot, Belg. vol. 1). 
His most important paper is the ^^Monographic de la Classe des 
FougSres," which is not a monograph but a paper on fern 
classification that is little known today. However, Bommer's 
classification was essentially that adopted by Eichler and later by 
Engler, and is thus quite similar to the classification adopted still 
later by Christensen, except for the inclusion of the Cyatheaceae 
in the Polypodiaceae. He collaborated with the Swiss botanist 
H. Pittier in publishing on the numerous then recent collections 
in Costa Rica of Pittier, Tonduz, Biolley, and others, in Durand 
and Pittier's 'Trimitiae Florae Costaricensis.'^ At the time of his 
death February 19, 1895, he was working on a monograph of the 
genus Adiantum. According to Dr. Lawalr^e, the manuscript has 
disappeared. Strangely enough, however, a synopsis of the pro- 
posed classification, in Bommer's own hand, is in the Smithsonian 
Institution, a Xerox copy of which has now been deposited in 
Brussels. Bommer can not be accounted an outstanding pteridol- 
ogist, but his influence is clearly evident in the Brussels herbarium, 
where his annotations are on many sheets, and where many of the 
older specimens in the general herbarium are still arranged as he 
left them,3 

Among the valuable parts of the old fern collections is the fern 
herbarium of Peter Claussen, but it seems that little is known of 
Claussen or his collections. It may be surmised that Linden and 
Claussen knew each other, since they were in Brazil at the same 
time, Galeotti must have been close to Linden, and through him 
with Funck and Schlim, Triana, and Lindig, for all these collections 
were turned over to Galeotti for the garden and for distribution. 
Thus Brussels is very rich in these old collections, especially those 

' For a bioeraohv. see lAo Errera. Bull. Soc, Roy. Bot. Belg. 34: 7-2L 1895. 

72 A^[ERICAN Fern Journai 

from Venezuela and Colombia that formed the basis of G. JVIetten- 
ius' treatment of the ferns in Triana and Planchon's "Prodromus 
Florae Novo-Granatensis" (Ann. Sci. Nat. [Paris] V, 2: 192-271. 
1864). The specimens in Brussels with the names in the hand of 
Mettenius should be considered the holotypes. These collections 
are partly represented by duplicates or fragments in the ]\Iet- 
tcnius Herbarium in Berlin. 

The Linden ferns are of particular importance. Jean Jules 
Linden was born in Luxemburg in 1817. At the age of IS he was 
authorized by the Belgian government to collect plants in tropical 
America with Nicholas Funck and Auguste Ghiesbreght as 
companions. They first went to southern Brazil. On the second 
trip Linden went alone to Cuba in 1837, and in 1838 to ^Mexico 
and Guatemala, where he obtained sizeable collections, especially 
in Tabasco and Chiapas. His third and most profitable trip was 
made with Louis Joseph Schlim in 1841 to Venezuela and Colom- 
bia, where he remained until 1844. In 1844-1845, Linden collected 
extensively in Cuba, especially in the eastern province of Oriente. 
On his return he established a nursery in Luxemburg in 1845, 
which was moved in 1851 to Brussels, where it became one of the 
foremost in Europe. In 1852 or 1853 he was appointed Director of 
the Jardin Royal de Zoologie et d'Horticulture de Bruxelles. 
Linden listed his offerings in catalogues that are rare and nmch 
prized today because they contain the original descriptions of 
some of the plants that Linden introduced to cultivation. In 1870 
he expanded his estabHshment by buying the large nursery of 
Ambroise Verschaffelt in Ghent. Linden's collections were never 
completely worked up, although some groups were studied, e.g. 
the "Orchidaceae Lindenianae" by J. Lindley. Linden sold his 
plants widely. There is a large set in Brussels, and an unexpectedly 
large set in the Webb Herbarium in Florence. Linden remained 
active until the age of 80, when he died in 1898, at the time the 
Consul-General in Belgium for Colombia. There is an account 
of his itineraries in Annales Soc. Roy. Agr. Bot. Gard. 2 : 205-220, 

259-272. 1846. 


Ferns in Euhopean Herbaria 73 

in Luxemburg. He was sent to ]Mexico with Linden in 1835 to 
collect, and in 1845 he went with Schlini to Venezuela and Colom- 
bia, where he assembled large and valuable collections. Un- 
fortunately many of these were lost in a shipwreck, but even so 
there are some 4,650 numbers of his collecting in Brussels. 

A collection of ferns of great importance is the Indian and 
Malaysian collection of William Roxburgh. I expect to publish a 
paper on the Roxburgh ferns, and so I shall not comment on them 
here other than to say that the Brussels set is the most authentic, 
with the names in Roxburgh's own hand. 

The Costa Rican ferns of Pittier, Tonduz, Biolley, Werckle, and 
others were worked up by Bommer, and later by the Swiss botanist 
Hermann Christ. Many of the Christ types from Costa Rica are 
in Brussels rather than in the Christ Herbarium, now in Paris. 

It is evident that when J. G. Baker was working on his account 
of the Cyatheaceae and Polypodiaceae for INIartius "Flora Brasil- 
iensis" (vol. 1, no. 2. 1870), that he had on loan the Brazilian 
specimens from Martius' herbarium, for they are annotated by 
Baker and correspond with the names as used in the Flora. In the 
case of the new species therefore Baker's types are in the Martms 
Herbarium in Brussels and not at Kew, where Baker's other 
types mostly arc. Sometimes Baker removed fragments and these 
are at Kew. The types of Johann Wilhelm Sturm (1808-1805), 
who wrote the accounts of the Hymnophyllaceae, Gleicheniaceae, 
jNlarattiaceae, Ophioglossaceae, Osmundaceae, and Schizaeaceae 
for the "Flora Brasiliensis," are also in the .Alartius Herbarium in 


The holotypes of the species described in 1^24 by H. A. Schrader 
from the Brazilian collections of the Prince von Xeuwied are also 
in the Herbarium. 

Martius lent his Brazilian collections to Gustav Ivunze, the 
noted pteridologist of Leipzig, who published his paper "Filices 
in ]Martii Herbarium Florae Brasiliensis" in 1839. These specimens 
were returned to Martius, and thus the holotypes of these Kunze 
species are still in existence, with the names in Kunze's hand, 
fortunately, since Kunze's own herbarium was destroyed in Leip- 

^^ AiMERicAN Fern Journal 

World War 

scribed by Kunze have also authentic material named by Kunze 

in Brussels. 

The other fern collections include those of Pohl, Karwinski, 
Blanchet, Luschnath, Glaziou (many numbers), Husnot, Gay, 
Bertero, and others far too numerous to mention. Naturally, 
Brussels is the richest herbarium for Belgian ferns, and those of 
France are well represented, due especially to the fine herbarium 
of J. Call6 and his collaborator E. Walter. 

Brussels is now fortunate in having pteridologists again for the 
first time since the late nineteenth century. The Director. F. 
Demaret, is interested in the ferns of the Congo. Dr. A. Taton 
also has studied African ferns and has published an account of 
the Congo Hymenophyllaceae. Dr. Andr6 Lawalr^e, the present 
pteridologist, prepared a perceptive account of the ferns for the 

Flore Gdn^rale de Belgique" (1950) and is now at work on the 
terns of the Congo, of which six parts have been published in the 

I^ ore du Congo, du Rwanda et du Burundi," the Equisetaceae, 
Fsilotaceae, Actiniopteridaceae, and Parkeriaceae in 1969, the 
bchizaeaceae in 1970, and the Blechnaceae in 1971. Collaborators 
preparmg manuscripts include Kramer (Lindsaeaceae), Launert 
(Marsileaceae), Bizarri (Selaginellaceae), and Pichi-Sermolli. 

I have visited Brussels three times, the first in 1954 to study and 
photograph the types of Martens and Galeotti. Prior to 1842, 
when their "JM^moire sur les Fougeres du Mexique" was published 
by the Acad^mie Royale de Bruxelles, very few ferns of Mexico 
were known. Martens and Galeotti described many new species, 
and they could have described more, for many that they identified 
with older species of the West Indies or South America proved 
to be wrongly identified and actually undescribed at the time. 
However, sometimes the Martens and Galeotti species belong to 
very cntical groups and they can be known definitely only by 
tne study of the original specimens. Many of the new species have 

ZJ^l '°''''''''"' ^""^ °^^"^^ ^^^^ Perhaps never been found 
Rn,L r! -f'^ °'^''^' duplicates of the Galeotti collections in 

Ferns in European Herbaria 75 

In my visits to Brussels I have always been most hospitably 
received, in 1954 by Dr. Robyns and Dr. Lawalrde, and in 1969 
and 1970 by Dr. Demaret and again Dr. Lawalr^e. Unfortunately, 
the ferns along with the general herbarium, including the Martius 
Herbarium, are now kept in a basement that is not well illuminated 
and possibly at times too damp. It is to be hoped that they can be 
moved soon to the new herbarium building that is ready for them. 

Much of my information comes from the paper by J. E. Bommer 
"Notice sur le Jardin Botanique de Bruxelles," which was written 
at the time when the present garden was first formed (Bull. Soc. 
Roy. Bot. Belg. 9: 418-455. 1S71). Some of the recent information 
comes from the publication that was issued last year for the 
centenary "Le Jardin Botanique National de Belgique, 1870- 
1970," which is unsigned, but is by Dr. Demaret, Dr. Lawalr^e, 

and their associates. 

(To be continued) 



Albert S. Rouffa* 

Over forty years ago Okabe (1929) wrote that . . . ''Psiiotum 
nudum, which grows wild in southern Japan, was once much 
cultivated in Japanese gardens as an ornamental plant. Slightly 
over 100 garden varieties were given fantastic names. Today, 
however, there are unfortunately only a few of them found oc- 
casionally in greenhouses of botanical gardens or in private 
gardens." Since the time that Okabe wrote this apparent epitaph 
there have been several attempts by Psilotum growing associations 
in Japan to revive the almost lost art and to gather information 
on the remaining varieties available. According to a special 

» Supported by a grant from the University of IlUnois, Chicago Circle 
Research Board. 

American Fern Journal 

Volume 61, Plate 14 

Pig, 1. A distal portion of an aerl\l shoot with five hranches of 



Appexdageless Psilotum 77 

issue in December 1967 of '^Horticultural Magazine" (Japan) en- 
titled ''Classical Horticultural Plants'' there are about 50 clearly 
identified old varieties of Psilotum plus about 10 unidentified 
and newly introduced varieties in existence. 

Among the older clones which may have been grown for more 
than 200 years there is one known as set ryu kaku, a descriptive 
appellation that may be broadly interpreted in the traditional 
sense as ''Green Dragon's Horns" or in a more modern sense as 
"Green Giraffe's Horns" (Rouffa, 1968). With all apologies to 
Okabe, the name, far from fantastic in this instance, appears 
rather appropriate (see Figs. 1-8). It is interesting to note that 
this Psilotum does not produce either sterile or fertile appendages 
and develops meiotic synangia at the tips of the ultimate branches. 
For this reason, this Psilotum merits an introduction and further 

Ihe clone was introduced into the United States in 1956 by 
Dr. John Creech (1957), of the New Crops Research Branch, 
U. S. Department of Agriculture, Beltsville, ^Maryland as PJ. 
235301 and, along with a number of other Japanese Psilotums, 
deposited and grown for a number of years at Longwood Gardens. 
Subsequently, some of the collection (not including this clone) 
was sent to the Xew York Botanical Garden. I am particularly 
indebted to Dr. Donald Huttleston of Longwood Gardens for 
the special propagation and donation of their Psilotums to our 
department in November 1967. Up to that time the unusual 
characteristics of sei ryu kaku were quite unknown to any of us 
until the plants were unpacked in our laboratory. 

I am also especially grateful to IVIr. Kaname Kato, an eminent 
horticulturist in Tokyo, for providing me with valuable informa- 
tion regarding this particular clone and the history and culture 
of Japanese Psilotums in general. 

Materials and Methods 

Severed shoot tips were treated in a number of ways for observa- 
tion with the Cambridge :Mark II stereoscan SEM (see figure 
legends) and included preservation in both glutaraldehyde and 

American Fern Journal 

VoLUMK 61, Plate 15 

Fig. 3. A distal portion of an aerial shoot with a sbries of branches 

VARIOUS lengths: Twn r i.^KTO_a,.,., ^^ 



P_ - A. """ xr.x.«iINATlNG IN SYNANGIA, X 2.0. 

AERt;; .Ir^^C^^^n r^^""^ ""^ ' '''^^^'^ ^^ SVNANGIA-BEAHING AND STERILE 

Append AGELESS Psilotum 79 

in FA A. A study is continuing to determine the best treatment 
for various conditions. Fresh tips could be observed for periods 
up to thirty minutes, when vacuum-induced shrinkage became 

Psilotums have been grown in our laboratories for a number of 
years indoors under fluorescent lights. They appear to grow best 
and produce more sporangia when cool-white lamps are supple- 
mented with wide-spectrum Gro-Lux lamps in a ratio of 2:1. Light 
intensities range from 1,000-2,500 ft-c. Sei ryu kakii has responded 
well to this lighting and grows as vigorously as many of the typical 
Psilotums. Despite the nearly constant cultural conditions, all 


of quiescence. 



appear relatively smooth and terete. Their often apiculate apex 
is unobscured by leaf primordia. Therefore one can easily observe 
the subdividing of the apex into multiple apices (Bierhorst, 1954) 
often Avhen the shoot is not more than 2 cm in length above the 
soil (Figs, 10, 11, 13), After its formation, each apex eventually 
develops its own subtending axis before the final elongation of 
the total aerial shoot takes place {Fig. 9, 11). The shoot apices 
continue to subdivide following elongation of the total shoot 
{Figs, 13, 14), The early subdividing of the emerging aerial shoot 
apex is synchronous and symmetrical {Fig. 13), but the final sub- 
divisions of the apices appear to be less synchronized and some 
may be quite asymmetrical {Figs. 14, 15). This asymmetry is 
perpetuated in the growth of some of the distal branches. As a 
result, distal shoots may develop monopodially and produce 
lateral branches ranging in size from mere ridges, to stub-branches 
1 mm in length, to nearly equivalent pseudodichotomies {Figs. 
1-5). Equal pseudodichotomies may also develop {Fig. 9), Unequal 
branches are often, but not always, the hallmark of shoots forming 
synangia {Figs. 1-8). 

American Fern Journal 

Volume 61, Plate 16 


Fig. 5. Shoot terminating in sterile forked-tipped uRANCii and a 

SYNANGIAL branch; glistening dots on SURFACE ABE STOMATE SITES, X 1. 

Figs. 6-8. Shoots terminating in short synangia-bearing pseudodicho- 

TOMiEs. Figs. 7 A 8. Saj 



Figs. 7, 8, X 1.5. Fig. 9. Actively elongating vegetative upper shoots; 


'1 absence of appendages, X 1. Fig. 10. Basal part of shoots near 
ground level; note absence of sterile-appendages and the naked 

Appendageless Psilotuai 81 

The stub-branches are suggestive of the sterile appendages of 
the typical leafy form of Psilotum, On the whole, shoot vascu- 
larization of sei ryu kaku appears similar to that of the typical 
form. Freehand sections taken at suitable intervals along the 
shoot and stained with HCl-Phloroglucinol reveal that the stub- 
shoots have a lignified strand branching from the stele extending 
at least to their base. Those stub-shoots bearing synangia are 
supplied with a lignified strand to the synangium. In certain 
respects, particularly in their vascularization, the sterile stub- 
shoots resemble the sterile appendages of Psilotum complamtum 
(Stiles, 1910). However, the stub-shoots are not necessarily pro- 
duced in helical order, although occasional distal shoots may 
show this arrangement {Fig. 3). 

Synangia typically develop at the tip of certain distal aerial 
branches. Such branches may be of varying lengths (F/f/s. 1-8). 
Meiosis has been observed in the synangia, and apparently normal 
kidney-shaped spores are eventually produced in the locules. 
The number of locules may be 4 or more rather than the usual 3 
(Figs. 1-3, 8), As in typical Psilotums, not all aerial shoots * 
sporangia, and there is a tendency for sporangium-bearing shoots 
to occur in groups (Fig. 4). Spore production (on a per- aerial shoot 
basis) may be lower for sei ryu kaku than for the typical Psilotum 


SHOOT APICES, X 1.2. Fig. 11. Closeup of basal portion of shoot about 
4 MM from soil levkl after formation of multiple apices, the upper 
shoot system beginning to elongate, X 2.5. Fig. 12. Meiotic metaphase 



14. Group of subdividing shoot apices from a slightly later stage than 
Fig. 13, the axes beginning to elongate; note the apical 



quite unsymmetrical; fresh and unco 

elongating shoots at a somewhat later stage of growth than Fig. 13, 


'^2 American Fern Journal 


where synangia are produced by a sei-ies of lateral fertile-append- 
ages along the distal ends of the aerial shoots. 

There is evidence of meiotic irregularities. Whether those 
observed were induced by environmental conditions under which 
the plants were grown (Okabe, 1929; Barber, 1957), or for other 
reasons, is not known. In initial chromosome counts of S:MC 
metaphases pretreated with oxyquinoline and stained with aceto- 
earmine, meiotic bivalents ranged from about 46-.>r) (e.g. Fig. 12). 
Sei ryu kaku is therefore a diploid. 


The entire scientific literature of Pstlotuni muhim aiid P. 
complanatum that deals with the origin and relationships of this 
genus has never considered any type of aerial shoot other than 
the one that produces reduced leaflike appendages of two inter- 
grading types: (i) a sterile series of awl-shaped appendages on 

shoot, ^.., „ _ ^,^_^„ „. .^...... 

appendages on the upper extremities of the shoot which is associ- 
ated with the development of synangia at each appendage (Bier- 
horst, 1956; Roth, 19G3a, b). In sei ryu haku, however, the lower 
extremities do not have leaflike appendages {Fig. 10). When the 
upper extremities of the shoot are differentiated they are also 
appendageless {Fig. 9), but stub-branches may be produced 
{Figs. 2, 3) by asymmetrical growth. These short stub-branches 
may resemble the sterile appendages of typical Psilotum, but 
are not produced in a continuous series nor is their number or 
size consistent. In this variety the synangia are borne only on 
the tips of branches {Figs. 1^8). 

The genus Psilotum and its companion genus Tmesipteris 
have long been considered modified subtropical and tropical 
rehcs of the early land plants of the lower Devonian. This was due 
m part to the apparent nature of their sporophytes, which seemed 
to bear little resemblance in growth and structural patterns to 
norQ °{^^*^'''' primitive plants. Recently, however, Bierhorst 
(19bSa, b, 1969) has taken another look at the problem and has 
shown a number of structural and developmental similarities 

Appexdageless 1*silotum S3 

between Tmesipteris especially and the enigmatic New Caledonian 
endemic fern Stromafopteris vioniliformis. Noting greater simi- 
larities between the two genera of the Psilotaceae and Stromatnp- 
teris than the latter possessed in regard to the family Gleichenia- 
ceae, to which Stromatopteris had originally been taxoiiomically 
assigned, Bierhorst (196Sa) proposed that this ai)parently 
anomalous fern be placed in a family of its own, the Stromatop- 
teridaceae. He further stated that the Psilotaceae and the 
Stromatopteridaceae were too closely related to be separated into 
separate orders. As a result he transferred Psilotum and Tmesipteris 
(Psilotaceae) into the order of ferns (Filicales). As a corollary, the 
aerial axes of Tmesipteris and Psilotum would then be considered 
homologous with the aerial "frond" of StrGViatopteris. Bierhorst 
(19G9) also postulated that the radially organized aerial axes of 
Psilotum, rather than being primitive, were actually specialized 
and derived from the winged, leaflike condition of Tmesipteris. 
These sweeping conclusions were based on an impressive amount 
of morphological and anatomical evidence, much of it gleaned 
from Bierhorst's own indefatigable collections in the field. 

The phytochemistry of the Psilotaceae has been recently 
enriched by the discovery of a phenolic glycoside, psilotin, that 
was isolated and identified from Psilotum nudum (:\lclnnes, 
Yoshida & Towers, 196;")) and Tmesipteris tannensis (Tse & Towers, 
1967). According to Tse & Towers the compound has not been 
identified in representatives of the Lycopodiophyta. 

There is a possibility that the phylogeny of the Psilotaceae, 
long dormant as a subject for scientific inquiry, may well agam be- 
come a subject of lively controversy. 

It may be that, in light of Bierhorst's conception that Psilotum 
has a derived radial shoot or axis, the unusual characteristics of 
sei ryu kaku may or may not have a direct bearing on the determi- 
nation or confirmation of the phyletic affinities of Psilotum. This 
clone is of more immediate morphogenetic interest because it 
performs routinely the production of terminal synangia, a phenom- 
enon that occurs only quite rarely, or under environmental stress, 
in most Psilotum genotypes. In typical Psilotum, spontaneous 

84 American Fern Journal 

proliferation of the fertile appendage into a csliort brauohlike struc- 
ture \va;s reported by Sykes (190S) and Bierhorst (1952). Typical 
Psilotiim grown under normal or greenhouse conditions have 
produced synangia in apparently terminal positions (Fabbri, 
1963; Abraham & Ninan, 1965; Rouffa, unpublished observations) . 
Also, Bierhorst (personal communication) has noted the existence 
of a native New Zealand clone in which the synangia are borne at 
the tips of distinct branches. Such occurrences could certainly 
strengthen the theory that the forked fertile appendage was 
evolved from a reduced branch or axial system with terminally 
produced sporangia, (Bierhorst, 19r)6; Rouffa, 1967). 

Indeed the resemblance of the elongated synangium tipped 
shoots of sei ryu kaku to the aberrant, elongated fertile appendages 
tipped with synangia produced for at least six years in the author's 
laboratory on a typical variety of Psilotum is rather striking (see 
Rouffa, 1967, ,/?f7, 4). The latter aberrant fertile appendages were 
induced under conditions of low light intensities, red spectrum 
deficiency, and high temperatures. The effect, although pro- 
duced quite regularly on an annual basis, appeared unstable 
since there was a tendency for the same shoot to return to the 
formation of the normal fertile appendage-synangium structures, 
even though the inducing environment remained unchanged. To 
explain this, I postulated that typical Psilotum still carries in 
its genome an ancestral set of genes responsible for the timing 
sequence of meiosporangium development at the tips of certain 
branche*^, but that some form of gene repression (possibly genes 
acquired through polyploidy) ''modernizes" the expression of 
the bran<*h-sporangium combination to that of a fertile appendage- 
sporangium combination. I further suggested that, somehow, 
repressing genes themselves were themselves repressed tempor- 
arily at the time of the experiment by the extreme stress of an 
unusual enviromnent, thereby unmasking temporarily the an- 
cestral condition where sporangia are produced terminally on 
elongated shoots. 

Set ryu kaku, however, normally produces synangia on the 
ultimate tips of shoots (both long and short) and does not produce 

Appenuageless Psilotum 85 

ai)peiulages. According to the above hypothesis, the ''modern- 
izing" genes are either continually repressed, or they are not 
present in the genome in this clone. Future studies on chromosome 
irregularities in this clone might shed some additional light on 
these possibilities. 

It cannot be denied that this much prized Japanese exhibition 
PsiloHnn looks for all the world like something out of the Devonian. 
However, in the face of new evidence, we must exercise caution 
in assuming that this variety, or any other form of Psilotum^ is a 
direct or relic descendent from tliat period. Nevertheless, there 
is a distinct possibility that in sei ryu Icalu some genetic change, 
as yet unilluminated, has stripped away several hundreds of 
millions of years of evolutionary modification and there is a re- 
mote chance that it represents one of the last of a nearly extinct 
race of Psilotum, 

Finally, it may be that we now have been provided tlie oppor- 
tunity to learn from its relatively simple structure how the early 
kind plants solved their developmental problems. We are now 
gathering for this purpose an array of variant genotypes of PsiJoium 

which should prove most useful for genetic and morphogenetic 
analyses and for phytochemical studies. 


Abraham, A. and C. A. Ninan. 1965. Morphological and cytological notes on 

Psilotuni nudum (L.) Beauv. Caryologia: 18; 537-539. 

Barber, H. N. 1957. Polyploidy in the Psilotales. Proc. Linn. Soc. New- 
South Wales %2: 201-207, 

Bierhorst, D. W. 1952. The morphology, anatomy and the phylogeny of the 

Psilotaceae. Ph.D. Thesi^s, University of Minnesota. 

1954. The origin of branching in the aerial shoot of Pbilotum 

nudum. Virginia J. Sci. 5 : 72-78. 

— . 1956. ObservatiouiS on the aerial appendages in the Psilotaceae. 

Phytomorph. 6: 176-184. 

— . 1968a. On the Stromatopteridaceae (Fam. Nov.) and on the 

Psilotaceae. Phytomorph. 18 : 232-268. 

1968b. Observations on Schizaea and Actinostachys spp., in- 
cluding A, oligostach^'s, sp. nov. Amer. J. Bot. 55: 87-108. 

^<' A.MEuicAx Fern Journal 

BiKRHORST, D. W. 1969. On Stromatopteris and its ill defined organs. Amer. 

J. Bot. 56: 160-174. 
Crekch, J. L. 1957. Plant explorations. Ornamentals in southern Japan. U. S. 

Department of Agriculture, Agr. Res. Serv. 34-1 : 1-55. 
Fahijri, F. 1963. Terminal sporangia at the apices of branches in two plants 

of P.silotum nudum (L.) Beauv. A preliminary report. Giorn. Bot. 

Ital. 70: 591-595. 

McInnes, a. G., S. YosHiDA, and G. H. N. Toayers. 1965. A phenolic gycoside 

from Psiloturn nudum (L.) Griseb. Tetrahedron 21: 2939-2946. 

NixAX, C. A. 1956. Cytology of Psilotum nudum (L.) Beauv. (P. triquetrum 

Sw.). La Cellule: 57: 307-318. 

Okaue, S. 1929. Uber eine tetraploide Gartenrasse von Psilotum nudum, 

Palisot de Beauvois ( = P. triquetnun, Sw.) und die trii>olige Kern- 
teilung in ihren Sporenmutterzellen. Science Reports, Tohoku Im- 
perial University, IV, 4 : 373-379. 

RovFFA, A. S. 1967. Induced Rsilotum fertile-appendage aberrations. Mor- 

phogenetic and evolutionary implications. Can. J. Bot. 45 : 855-861. 
1968. An appendageless form of Psilotum. Amer. J. Bot. 55: 

714. (Abstr.) 

1963a. Histogenese der Luftsprosse und Bildung der "dichotomen" 
Verzweigungen von Psilotum nudum. Adv. Front. PI. Sci. 7: 157-179. 
"'■"■ Hist 

Psilotum nudum. Flora (Allgem. Bot. Zeit.) 153: 90 111. 
Stiles W. 1910. The structure of the aerial shoots of Psilotum flaccidum. 

Wall. Ann. Bot. 24 (O.S.) : 373-387. 
Sykes, W. G. 1908. A note on an al>nornialitv found in Psilotum triquetrum. 

Ann. Bot. 22 (O.S.) : 525-526. 
TsE, AiDA and G. H. N. Towers. 1967. The occurrence of psilotin in Tmesip- 

teris. Phytochemistry 6: 149. 


AT Chicago Circle, Box 4348, Chicago, Illinois 00680. 

\'a8CULarization of Fekx Leaves S7 

Vascularization of Fern Leaves 

Robert C. Lommasson and C H. VounGj Jh. 

The subjocl of fern leaf venation is a very old topic and it 
has been dealt with by many authors during the nineteenth and 
twentieth centuries. Most of the discussions of fern leaf venation 
concentrate on its qualitative aspects resulting in descriptions of 
\'arious types of leaf venation. Considerably less attention has 
been devoted to the quantitative aspects of fern leaf vascular- 
ization. This report will give a brief summary of some of the 
measurable aspects of the vascular system within the leaves of 

certain rather common ferns. 

The collections of the ferns used in this study came from two 
sources. Those collected out-of-doors came from Giant City State 
Park or from Jackson Hollow, six miles northeast of Simpson, 
Illinois. ALitcrial was taken with the permission and under the 
supervision of Dr. Kobert H. Mohlenbrock of Southern Illinois 
University. Other ferns were obtained from the Department of 
Botany greenhouse at the University of Nebraska. 

The fern leaves were studied from slides made of cross sections 
of the material which had been stained with Delafield's hematoxy- 
lin and safranin. Whole mounts of cleared leaves or leaflets or parts 
of these were also studied to obtain vein patterns and angle of 


All measurements recorded in this study were made by means 
of an ocular micrometer. All those listed in Table I are averages 
of many separate measurements. Leaf thickness was measured 
at veins rather than between veins. Intervascailar interval was 
measured from the vascular tissue of one bundle to the vascular 
tissue of the next adjacent bundle; thus it includes any specialized 
parenchyma or bundle sheath cells. Vein width includes only the 
width of the conductive tissues of a vas(ailar bundle. Angle of 
branching refers to the branching of vascular bundles from the 
midvein of the leaf or segment and not to the ultimate branchings 
in the lamina. It was measured in linear units and the angle 

SS American Ferx Journal 

computed as a trigonometric function. Dissection of leaves as 
used in this study was an arbitrary rating scale which divided 

Table I. Summary of Leaf Measuremknt8 of Some Common Fekns 


Species Source in /t in fi in /x degrees D 

Bolrychiuin virginianum I 260 747 85 39 5 

Osmiinda cinnamomea I 125 426 32 41 3 

Trichomanes boschianum I 98 799 45 51 1 

Pteris vittata G 108 463 53 43 2 

Pteris cretica G 174 791 79 39 2 

Cheilanthes lanom I 135 281 28 43 4 

Pellaea atropurpurea G 195 396 45 41 4 

Adiantum pedatum G 56 622 38 48 o 

Adiantum capillus-veneris G 72 323 20 32 5 

Nephrolepis exaltaia G 176 561 45 38 2 

Woodsia obtusa I 177 429 39 37 3 

Pohjstichum acrostichoides 1 286 496 44 53 2 

Cyrtomiiim falcatum G 336 744 78 42 2 

Dryopteris dentota G 108 404 43 42 3 

Dryopferis marginalis I 171 607 42 42 4 

Dryopteris spin u losa I 110 633 38 40 6 

Dryopteris hexagonoptera I 99 581 18 45 3 

. 1 Ih yrivm fdix-fem ina I 1 38 437 30 33 4 

Asplenium trichomanes I 90 429 30 34 2 

Asplenixim platynenron 1 140 500 38 35 2 

Caynpiosorns rhizophyllns I 219 984 57 62 

Pohjpodium vtdgare G 176 736 78 43 1 

Polypodium virginianum I 200 681 42 44 1 

Phlebodiiim mtrenm G 1141214 69 44 1 

I = Southern Illinois; G = Greenhouse, Univ. of Nebr.; LT = leaf thickness; 
IVI = Intervascular interval; VW = vein width; AB = angle of branching; 
D = Disiiection of leaf: = entire; 1 = pinnatifid; 2 = pinnate; 3 = pin- 
nate-pinnatifid; 4 = twice-pinnate; 5 = more than twire-pinnate. 

fern leaves into six groups. These were designated by a numerical 

those which were more than twice pinnate. 


Vasculauization of Fern Leaves S9 

Leaf Thickness 

111 general, the leaf thickness at vascuhir bundles is greater 
than it is between them. In five species the leaf thickness at the 
vascular bundle exceeded the sum of leaf thickness between veins 
and the vein width. This indicates an actual build-up of tissue 
around the vein. These five species are Osiminda cinnamomea, 
Trichoiiianes hoscliianum, Woochia ohliisa, Dryopieris hexagonop- 
tera, and Asplenium plahjneuron. The leaf blade is thicker be- 
tween veins than it is at the veins in three species, namely Pteris 

rittata, AcUanlum capillus-venerisy and Dryoptens marginalis. The 
measurements listed for LT in Table I are averages of those taken 
at the vascular bundles. 

There appears to be no great modification of leaf thickness 
due to environment as judged from leaves collected from the two 
sources. The averages of leaf thickness for these fern leaves was 
from 50 microns in Adianlum pedatum to 336 microns in Cyrtcmiwn 

Intervascular Interval 

IMeasurements of Wl were taken where the adjacent vascular 
bundles were nearly parallel- Tlie predominantly dichotomoas 
branching of lateral vascular bundles sometimes occurs near the 
midvein as in Osmunda cinnamomca, Dryopieris hexagonopiera, 
and NephroJepis exaUata. In Plen's vitlata some lateral vascular 
bundles branch near the midvoin and adjacent ones branch mid- 
way to the margin or very near the margin. Other ferns, such as 
Adiardum pedatum and Aspleiihim pJalyneiinm, show several 
branchings of one vascular bundle. 

Ferns ^^ith the largest leaf segments have correspondingly 
lai'ge intervascular intervals. Table I sliows that for Phlebadmm 
aiireum, Camptosorus rhizophyllus, Plen's crcticay and Cyrtommm 
falcatum the IVI is over 700 microns. Other ferns having inter- 
A-ascular intervals in this range, however, include plants with 
smaller leaf segments, such as Boirychium virginiamm, Tricho- 
manes hoschianum, and Poly podium vidyare. 

90 Ameuican Fern Journal 

Comparisons of IVI with the dissection of leaf (D) Hsted in the 
right hand column of Table I have been illustrated in Fit/. J. 
Except for group 5, the IVI measurements for each group de- 
crease as the dissection of the leaf increases. In Fig. 1 the wide 
variations within each group are not related to the source. For 
example, the greatest IVI in group 1, Phlehodium aiireiim, and the 
smallest IVI in group 5, Adiantum capillus-veneris, were both 
grown in the greenhouse. It might be profitable to study the 
combination of segment size and leaf dissection as it may be 
related to intervascvdar interval. 

Vein Width 

Vein width measurements show no constancy or correlation 
with source of material, genus, leaflet size, or leaf dissection. The 
range of vein width from 18 to 85 microns indicates that the tissue 
requirements of each species govern the size of veins. 

Angle of Vein Branching 

In the genera Dnjopteris and Polypodhim the species studied 
show similar angles of branching of the veins. In Woodsia and 
Adiantum the two species of each genus which were examined have 
considerably different angles of branching. Woodsia obtusa shows 
37° and Woodsia scopulina 30^ (not shown in the table). The 
Adiantums are also very different, with angles of 32^ and 48°, 

Since the angle of branching is measured from the midvein 
of the segment, the degree of dissection probably affects the angle 
at which lateral veins arise. The midveins in a highly dissected 
leaf are farther removed from the rachis. A correlation of the 
degree of dissection and the angle of branching, however, failed 
to show any significance for the species studied. A more detailed 
and refined treatment of these characteristics might be profitable. 


The tissues of the fern leaf do not show some of the specializa- 
tions commonly seen in the leaves of higher plants. Tlie epidermis 
is not as highly specialized unless the inner lobes filled with ehloro- 
plasts are considered to be a specialization. The mesophyll of 

Vascularization of Ferx Leaves 


many ferns is not noticeably differentiated into palisade and spongy 
layers as are many dicotyledonous leaves. This condition empha- 
sizes the horizontal continuity of the leaf tissues. The build-up of 
tissues around the veins of many ferns also allows for the greater 
distributive and collective area for the vascular bundle, as em- 
phasized by Armacost (1944). 


In u 






































2 2 2 2 ? 2 2 

3 3 5 3 

4 4 <^ ^ 

5 5 5 

Fig. 1. A histogram of certain" coMMO>f ferns arranged in groups 


Although the average leaf thickness in this study of fern leaves 
is 149 microns, about one half of the species have a thickness 
between 60 and 140 microns. This is strikingly similar to the 
findings of Wylie (194()) in his study of ninety species of Florida 
dicotyledons in which 53 species had an average thickness ■withm 
this range, but the average thickness of all 90 species was a6 

92 American Fern Journal 



study (Wylie, 1939) of 22 Iowa woody plants and 24 Iowa her- 
baceous plants J which had leaves averaging 152 and 163 microns 
respectively, these ferns are not significantly thinner. The factor 
which gives the impression of thinness in fern leaves is the absence 
of the abundant network of major veins that gives structural 
rigidity to many dicotyledonous leaves. 

The vein width has already been referred to in estimating the 
build-up of tissues around the vein. Although the veins in ferns 
are somewhat smaller at their tips and larger at the base of leaflets, 
many ferns fail to show, aside from a large midrib, the degree 
of difference referred to in other types of leaves as primary, second- 
ary, tertiary and so forth. This can be explained by one of the 
principles of Eower (1923) which says that in most fern leaves 
there persists a distal dichotomy in the venation. Ihe vein width 
does not correlate with the I VI, the angle of branching, or the 
degree of dissection of the leaves. It does correlate weakly with 
leaf thickness. Eower (1923) also suggested that in evolution the 
factors of leaf venation and leaf dissection do not advance together. 

If the method of Ettingshausen (18G1) in measuring the angle 
of branching had been used, more consistent results might have 
been obtained and the angles would have been larger. 

Wylie (1939) mathematically defined the relationship between 
tissue organization and vein distribution in dicotyledonous leaves. 
He showed that with large ratios of spongy to palisade mesophyll 
the intervascular interval was large, and conversely, when there 
was a relatively thick palisade layer the intervascular interval 
was correspondingly small. Later Wylie (1948) pointed out the 
dominant role of the epidermis in two species of Adianlum, The 
lobed epidermal cells of ferns can be considered spongy mesophyll 
since they are parts of a horizontally contiguous tissue. In sonie 
ferns there is little or no mesophyll which may be considered 
to be vertically oriented palisa^Ie mesophyll. We should not be 
surprised, therefore, to find such large intervascular intervals in 
the ferns as compared with angiosperm leaves. In this study the 
average of all IVI measurements was 595 microns. For 194 species 

Vascularization" of Fern Leaves 93 

of dicotyledons from Iowa, Florida, the Pacific coast, and Xcw 



average I VI of 155 microns. Lommasson (194,S) in a study of 37 
species of grasses found an average lYl of 141 microns. Since the 
IVI represents the distribution of the smallest veins of the leaf 
and since they are among the last tissues of the leaf to develop, 
they represent the final adjustment of the vascular tissue to the 
environmental conditions in which the plant is growing. These 
results again point to the often suggested idea that vein spacing is 
dependent upon the tissue organization of the leaf and that this in 
turn is governed by the demands of the constituent chlorophyll 
bearing cells. 

Literature Cited 


and vein-rib of certain dicotyledon leaver. Proc. Iowa Acad. Sci. 

51: 157-169. 
Bower, F. O. 1923. The Ferns. Vol. 1. Cambridge. 
Ettixgshausen, C. R. von. 1861. Die Blatt-Skelete der Dikotyledonen. 

LoMMAssoN, R. C. 1948. Tissue relations of grass leaves. University of Iowa, 

Phu.pott, Jane, 1953. A blade tissue study of leaves of forty-seven species of 

Ficus. Hot. Gaz. 115: 15-35. 

Wyije, R. B. 1939. Relations betw^een tissue organization and vein distribu- 
tion in dicotyledon leaves. Amer. J. Bot. 26: 219-225. 

' — . 1946. Relations between tissue organization and vascularization 

in leaves of certain tropical and subtropical dicotyledons. Amer. J. 
Bo^ 33: 721-726. 

" . 1948. The dominant role of the epidermis in leaves of Adiantum. 

Amer. J. Bot. 35 : 465-473. 

— . 1954. Leaf organization of some 
Zealand. Amer. J. Bot. 41: 186-191. 

University of Nebraska, Lincoln, Nebraska 6850S. 

l)i American Fern Journal 

Shorter Note 


A Wkonglv Localized Species of PYnnoaiA.—Niphobolus 
cuneatus Kuhn (Bot. Zeit. 26 : 40. 186S) was described from material 
distributed by Hohenaeker "in territorio rei publ. Ecuador" no. 5. 
Since Niphobolus was considered a synonym of Cyclophorus by 
Christensen, the new combination Cydoplioms cuneatus (Kuhn) 
C. Chr. (Ind. Fil. 198. 1905) was pubHshed, but apparently no 
pteridologist has ever looked carefully at the type since the 
original pubUcation of the species. 

Both Niphobolus and Cydoplioms are later names for the genus 
now known as Pyrrosia. In the "Index Filicum," there were only 
two American species recognized, both of them a query— C. 
amencanus (Hook.) C Chr. and C. cuneatus. The former was 
properly considered a different genus, Niphidium, by John Smith. 
And thus C. cuneatus remains as nominally the only American 
species of this genus. 

Through the courtesy of Dr. Eekhardt and Dr. ]Meyer, the type 
of .V. cuneatus has been made available to me on loan. It is a 
minute specimen, containing about a half dozen fronds and a 
short length of rhizome. The material is sterile only. However, 
even in the absence of fertile fronds, it is readily identifiable as 
PyrrosJa rupestris (R. Brown) Ching, a species confined to Aus- 
tralia. The minute, obovate sterile blades, densely covered be- 
neath with stellate scales, these with a conspicuous, minute, round, 
red central disk and elongate straight white rays are characteristic; 
the rhizome scales agree also, these being elongate-lanceolate, 
light brown, and not ciliate. 

It is not at all likely that this Australian species occurs in 


distributed plants from many parts of tlie world and a confusion 
of data is not at all improbable. The genus Pyrrosia is therefore 
to be excluded from the flora of the Xew World.— C. V. Mouton, 
X at loml Museum of Natural History. Washinntnn nn m^m. 

IvECENT Vkhs Literature 95 

Recent Fern Literature 


The Genus Adiantum l\ Cultivation [Polypodiaceae], by 
Barbara Joe Ho^hizaki. Baileya 17, 110s. 3-4, pp. 97 -19L/. /— ,^7. 
197L Published by the L. H. Bailey Hortorium, Cornell University, 
Ithaca, N. Y., SI. 50. — In continuing her studies of cultivated 
ferns, Barbara Joe has given us a useful treatment of the maiden- 
hair ferns, which are popular with florists. These beautiful ferns are 
delicate, and do not like the hot dry atmosphere of most houses 
and apartments. Consecjuently, they are best grown in green- 
houses, where most of them are of relatively easy culture. Twenty 
_)ecies are recognized as being in cultivation in the United States, 
but almost all the named cultivars belong to Adiantum raddianum 
(usually called A, ciineatum), the ''Delta [Maidenhair." The text 
is made most usable by the good line drawings of typical pinnules 
and by silhouettes of the blades. Many of the named forms are 
not easily determinable, because they vary greatly according to 
cultural conditions and also intergrade with each other extensively. 
It is indicated that some of the cultivars can arise repeatedly 
de novo from normal parents. This is evidently a genus that would 
repay genetical study.^— C.V.M. 

Pollen and Spokes of Chile, by Calvin J. Heusser. Pp. 1 — Qo, 
?>/. 1—60, 1971. Published by the University of Arizona Press, P. 0. 
Box 3398, College Station, Tucson, Arizona 85700, $15.00. The 
present work describes, keys, and illustrates by photomicrographs 
54 species of Pteridophyta of C liile, the Juan Fernandez Islands, 
and Easter Island [which is rather remote from Chile in its flora 
and more closely related floristically to Polynesia]. The work 
appears to be carefully done, using the Erdtman method of spore 
preparation. However, it mav be doubted that this approach is 
truly valid at present. The scanning electron microscope is rev- 
olutionizing the study of spores, for it shows up many details 

' ^Irs. Ho^hizaki hari a^^ked me to point out twu typographical errors: On 
p. 104, "2-3 piiHiate'' is transposed with "3-4 pinnate"; ou p. 132 "Aa" is a 
pinna, not a frond. 

9G Amehican Fern Jourxal 

that can not be seen at all with the ordinary niioroscope, even 
with great magnification. The present photographs are hardly 
good enough for a clear identification of the species. The only 
out-and-out error that I notice is the inclusion of the description 
and illustration of the spores of "Davallia valdmana," although 
there is no such published species. Since the specimen concerned 
is deposited in the museum in Santiago it could have been properly 
identified. The genus Davallia does not grow in Chile, and so I 
would guess, without evidence, that the plant concerned is Rumohra 


and although 

common in the Province of Valdivia is not otherwise included in 
this treatment. Since most of the text and illustrations in this 
work concern the phanerogams, the book will be of more use to 
students of the flowering plants.— C.V.AI. 


Flora de Venezuela, Vol. I. Helechos, by Volkmar Vareschi. 
Instituto Botanico, Aptdo. 2ir^C), Caracas, Venezuela. 1033 pp. 
19G8.— This volume, which is bound in two portions, illustrates 
well the difficulties of preparing detailed floras in tropical areas. 
Although about 700 species are treated, in some groups (e.g., 
Polypodium and the Cyatheaceae) the list of "critical species," 
those that are published but which were not studied by the author, 
contains 60-80% as many as those accounted fur in the flora! 
The volume is dated 1968 on the title page and 1969 on the soft- 
bound cover, but dates in the text indinate that the writing was 
completed as early as 1960. Critical recent literature is often not 
cited nor additions and corrections apparently made. Most sur- 
prismg is the seriously flawed nomenclature. About 200 names are 
invalid for one or more reasons, including the lack of a Latin 



13 uninten- 

tionat. Of the former, 2 are illegitimate and 29 are legitimate; 
of the latter, 6 are illegitimate and 7 are legitimate. Reference is 
made to specialized details of anatomy, morphology, and paleo- 
botany, which IS useful and missing from most floras. The illus- 
trations are well executed and helpful.— D B L 

ExDtie aod Hardf Ferns 



2131 Valleio Street 
Sh Heleno, Californio 94574 


963-299S— Area 



A Serwfice ta Biatagwsis 

To better serve tlie biological eommnnlty: 

1. TRIARCH produces and stocks nearly 3000 dif- 
ferent prepared microscope slide items for use in gen- 
eral botany, plant anatomy, plant physiology, phyto- 
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For our current catalog No. 15^ or to request special 

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RipoB, UlaeoKsiM S4971 


Vol. 61 July-September, 1971 No. 3 

American Jfern Journal 


Published by tht Ms S SOU Hi BoTAl^^iCTCtJ 


DEC fi - 1971 



_ i L.i3??ABY. 







The Fern Vegetation of Aldabra AtoU. . , F. R. Fosbbrg 97 

Leaf Epidermal Studies in Marsilea 

John T. Mickel and Frank V. Votava 101 

The American Species of Plagiogyria sect, Carinatae 

David B- Lellinger 110 

The Genus Stenochlaena J. Smith with Description of a New Species 


The Variation in Spore Size and Germination in Dryopteris Taxa 

Dean P. Whittier and W. H. Wagner, Jb- 123 

The Gametophytes of Nattu*al Hybrids in the Fern Genus Pellaea 

Thomas R. Prat 128 

Schemochromic Blue Leaf-surfaces of Selaginella 

Denis L, Fox and James R. Wells 137 

Shorter Notes: Asplenium kamtchatkanum Gilbert, a Dubious Eastern 
Asiatic Fern ; A New Locality for Psilotum nudum in Sonora, Mexico . 140 

Recent Fern Lterature Mt^^PV ?i .^.9Ttr^Jf A^ .... 142 

DEC 6 - 1971 

®l)e American Jfern ^ocietp 

Council fot 1971 

Warren H. Waqnkr, J 
Ann Arbor, Michiga 

John T, Mickel, New 
York. 10458. 


, Bronx, New 


Richard L. Hattke, Department of Botany, University of Rhode Island, 

Kingston, Rhode Island 02881 Secretary 

LeRov K. Henrt, Division of Plants, Carnegie Museum, Pittsburgh, Penn- 
sylvania 15213. , Treasurer 
David B. Lellinoer, Smithsonian Institution, Washington, D. C- 20560, 


j^ational ^ocictp ^eprcfientaHbcs 

Warren H. Wagner, Jr., University of Michigan A,AA.S. Representative 
RoLLA M. Trton, Jr., Harvard University AJ,B,S. Represenlattve 

iamcritan jFcrn journal 


David B, Lbli.inger Smithsonian Institution, Washington, D. C. 20560. 

C V. Morton Smithsonian Institution, Washington, D. G. 20560. 

RoLLA M. Tryon, Jr. 



An illustrated quarterly devoted to the general study of ferns, owned by 
the American Fern Society, and published at 3110 Elm Ave., Baltimore, 
Md. 21211. Second-class postage paid at Baltimore. The pages of the Jour- 
nal are open to members who wish to arrange exchanges; membership lists 
assist those interested in obtaining specimens from dilTerent localities. 

Matter for publication should be addressed to the Editor-in-Chief. 

Changes of address, applications for membership, subscriptions, orders 
for back numbers should be addressed to the Treasurer. 

Subscriptions $5.50 gross, $5.00 net (agency fee $0.50); sent free to mem- 
l>ers of the American Fern Society (annual dues, $5.00; sustaining membership, 
JIOJK); life meml^ership, $100.00). Extraete<i reprints, if ordered in advance, 
will be furnished authors at cost, plus postage. 

Back volumes $5.00 to $6.25 each; single back numbers of 64 pages or 

less, $1.25; 65-80 pages, $2.00 each; over 80 pages, $2.50 each; Cumulative 

Indtx to Volumes 1^25, 50 cents. Ten percent discount on ortiers of six volume 
or more. 

Hibcarp anti ilerbarium 

Dr. W. H. Wagner, Jr., Department of Botany, University of Michigan, 
Ann Arbor, Michigan 48104, is Librarian and Curator, Members may borrow 
books and specimens at any time, the borrower paying all shipping costs. 

S>j[wre Cxcljangt 

^ Mr. Neill D. Hall, 1225 Northeast 95th Street, Seattle, Wa8hingtori981 15, 
IB Director, Spores exchanged and collection lists sent on request. 

hvm an 

#ifts anft IPettticstfi 

nable it to expand 
Herbarium aneeim' 

k issues of the Jo0BNal, and cash ^. ,^^^ 

tax-deductible. Inquiries should be addressed to the Secretary. 



^meruan jFern fournal 

Vol. 61 July-September, 1971 No. 3 

The Fern Vegetation of Aldabra Atoll 


Aldabra is a ring-shaped coral island in the western Indian 
Ocean, 260 miles northwest of the northern tip of Madagascar 
and 400 miles east of Africa. Unlike the majority of coral atolls 
it has been elevated by tectonic forces about 5-6 m. The island is 
noted, for more than any other reason, as the last stronghold in 
the Old World of the giant tortoises that formerly roamed many 
regions of the earth. They now survive only in the Galapagos, 
where they are rare, and on Aldabra, where they are still abundant. 

The climate of Aldabra is dry and hot, and scarcely favors the 
development of much of a fern flora. Although ferns are prominent 
in several habitats, the fern flora is exceedingly poor. One species 
had been recorded prior to the Royal Society Expedition (1966- 
1967) and, despite some week? of active collecting by two botanists 
and several other scientists interested in the vegetation, the known 
fern flora still remained a single species, Acrostichiim aureum L., 
until a tiny colony of Nephrolepis hisenala (Swartz) Schott was 
found by INIr. S. A. Renvoise in a depression in a barren area of 
windbeaten rough limestone near the southeast coast. No further 
colonies, nor any other species, have been found on the atoU- 

That promipent and varied fern vegetation may be made up of 
essentially a single species is very odd, indeed. Acrostichum is a 

^ The observations reported here were made while the author was a member 
of the Royal Society Aldabra Expedition, in January and February, 1968. 
Thanks are due to the Royal Society and to the Smithsonian Institution's 
Office of International Aclivitieji for the privilege of participating in this 


Volume 61. No. 2 of the JurRNAL, dd. 49-96, was issued June 24, 1971. 

American Fern Journal 

Volume 61, Plate 17 

Sv^fr' h^^*^"^^ ''champignon'^ WITH ACROgTICHUM AND TORTOISE. FiG. 2. 

by Jay sSe?''"™ ^^^^^^ ^i^ants in coral pits nkar the beach. Photos, 

Ferx Vegetation of Aldabua Atoll 99 

big, coaise, leathery fern with erect fronds. It is usually associated 
with mangrove swamps and coastal marsh vegetation throughout 
most of the Tropics. On Aldabra there are notable mangrove 
swamps, but Acrostichum was not seen in any of them. Tu vis- 
ualize where it does occur a brief account of the main surface 
features of the island is necessary. 

The island is a flat, almost complete ring of reef limestone, little 
higher than 7 m except for dunes, broken by several channels 
through which the tides race twice a day in and out of the great, 
shallow lagoon. The land area is about GO square miles and, 
although entirely of limestone, exhibits several different types of 
surface. \Much of it is extremely eroded into sharp pinnacles, 
ragged projections, deep pits, fissures, chimneys and crevasses. 
The edges are razor-sharp where the relief is too high for the 
tortoises to climb over it {Fig. 1, background), but where the relief 
is more moderate, the rubbing of their great shells has rounded 
off the projections {Fig. 1, foreground). Ihis rough type of surface 
is locally called ^'champignon"; in Polynesia it is called ''feo." In 
the interior of some of the islands are areas of a f:at, smooth or 
often slab-covered surface, called ^'platin/' with solution basins, 
fat soil areas, deep cracks and fissures, and numerous pools of 
water and mud. The pools have either sloping or vertical sides, 
and the water varies from fresh to as saline as sea-water or even 
more so. Storm beaches and occasional dunes of coral sand line 
some of the coasts, as on the northeast, east, and parts of the south 
sides of South Island, which is by far the largest of the four main 
sections of the ring of land. ■ 

It is not surprising to find that luxuriant masses of Acrostichum 
entirely or partly fill many of the pools, especially and almost 
exclusively those with vertical sides and mud bottoms. It tolerates 
a remarkable range of salinities, by actual tests ranging from 1,700 
to 15,700 parts per million dissolved salts. In these pools the 
masses of thick, creeping, branched rhizomes are dense, and the 
erciit fronds arc crowded in compact clumps which reach 1.5 or 
even 2 m in height from the mud. The larger pools could appro- 
priately be called marshes- 

100 American Fern Journal 

One sizeable area which I visited has a number of such pools, 
filled with Acrostichum, but in addition, several hectares of "platin" 
surface are intricately cracked. The cracks are filled with Acro- 
stichum. The rhizomes are mostly in the tops of the deep but 
narrow cracks; the abundant roots go down to the water as much 
as one meter below. The fronds here are smaller than in the pools, 



L. make parts of this area look like a miniature fern "savanna." 

Even more remarkable was an extensive area 100-300 meters 
vdde of "champignon" extending along the east coast of the island 
back of the line of dunes {Fig. /). In most of the island the "cham- 
pignon" is covered by thick scrub, but here, possibly because of 
exposure most of the year to salt-spray blown by the tradewinds, 
there is little woody vegetation. Instead, in addition to a few herbs 
and dwarf shrubs hiding in fissures, pits, and chimneys, Acrosti- 
chum is the principal component of the vegetation (Fig. 2) . Deep 
pits, a few centimeters in diameter, occur in great numbers in 
this limestone. Many of these have each an erect tuft of dwarfed 
Acrostichum fronds rising from a bit of rhizome down in the bottom 
of the pit. The fronds are from a few cm to 0.5 m tall, and even 
taller where they grow in chimneys or crevasses that afford better 
shelter. Occasional tiny plants grow in deep crevices. The whole 
area IS very sparsely covered by these tufts of erect, green, leathery 
fronds. Many of the fronds have several terminal pinnae densely 
covered on the dorsal side by sporangia, seen under a lens to be 
arranged m a close network. Pits in rough limestone in almost any 
part of the island are likely to be occupied by Acrostichum, but 
along the east coast it is the principal vegetation on large areas. 

That one species of fern is able to assume such variations in 
nabit and to tolerate such a range of conditions that it can fill a 
series of niches normally occupied by other species shows the 
capabihties that may be brought out by a wide variety of habitats 
and little competition from other species. An interesting question 
wV if ^^'•^f ^^^^ is not found in Aldabra's mangrove swamps. 
Which usually are regarded as its normal habitat. Perhaps there 

Leaf Epidermal Studies in Marsilea 101 

IS merit m the suggestion that the mangrove fern is actually a 
species of Acrostichum other than A. aureum L. Watson (]Malayan 
For. Recs. 6: 152-137. 1928) has distinguished A. speciosum 
Willd, as a species tolerating more frequent tidal inundation. No 
plants corresponding to the description of this, which has con- 
spicuously acuminate pinna tips, have been found in Aldabra. 

Departiment of Botaxy, U. S. National ]\Iuseum, Wash- 
ington, D. C- 20560. 

Leaf Epidermal Studies in Marsilea 

John T. IMickel and Frank V. Votava^ 

The aquatic fern genus Marsilea is widespread and well known 
throughout the world. Of the nearly 70 extant species, 20 are 
found in the New World. ]\Iany studies have been made on the 
genus from the standpoints of its physiology, morphology, 
anatomy, and development. Frequently there are difficulties in 
species identification since the taxonomic characters lie primarily 
in the reproductive structures — the sporocarps — and herbarium 
specimens are often sterile. A search was begun for vegetative 
characters to distinguish the species of^North America, Mexico, 
and the West Indies, As part of this study leaf clearings were 
made, and some notable diversity was found within the group. 

Gupta (1957), in a preliminary study of the epidermal and soral 
characters of some American species of Marsilea, reported that 
the epidermis could be of some taxonomic Value. Species differ, 
he said, in the walls of the epidermal cells ("smooth, slightly wavy, 
and highly wavy") and in the frequency of the stomata. He 
illustrated the epidermal patterns of 11 species and counted and 

* We wish to acknowledge the support of the National Science Foundation 
(Grant no. GB-13722 to the senior author). 


A.^iERiCAN Fern Journal 

measured the stomata. His monograph of the Indian species 


(1962) reported differences in epidermal patterns under different 
ecological conditions. 


LOWER. Fig. 3. M. 

Fig. 1. M. quadrifolia, upper. Fig. 2. Same, 

iCANA, upper. Fig. 4. Same, lower. Fig. 5, M. poly- 
CARPA, UPPER. Fig. 6. M. naphii, upper. 

Our clearings of 13 species were studied in regard to stomatal 
size, frequency and position, epidermal patterns, and certain 
ecological effects. Our findings differ somewhat from and go 
beyond those of Gupta sufficiently to merit reporting them here. 

Clearings were made of all 13 species found in North America, 
Mexico, and the West Indies. The specimens are cited in Table I, 
and all are in the herbarium of The New York Botanical Garden. 
Leaflets were cleared in 5% KOH for 1-2 days and were then 

Leaf Epidermal Studies in Marsilea 103 

bleached in 20 % Chlorox solution. The clearings were stained with 
tannic acid and ferric chloride and mounted in diaphane. 

Epidermal patterns 



6, 7), and (ii) those with wavy walls that range from only slightly 
wavy to strongly undulating {Figs. 8-15) ; but we could distinguish 
another group, those with short, blocky cells {Figs, 1-5), 

Smooth walls are quite distinctive and are found in M. nashii 
and M. tenuijolia. This character may be related to their very 

slender pinnae. ■ 

Short, blocky epidermal cells were found consistently in our 
specimens of M. qiiadrifdia, M. pclycarpa^ and M. mexicana- Their 
walls vary in wall waviness, even within a species. In M. pclycarpa 
the cells tend to be more elongate than in the other two species. 

JNIost species have somewhat elongated cells with variously 
undulating walls {Figs. 8-15). There seems to be considerable 
variation, and we found it impossible to separate sharply those 


mitcronata. M 


been combined by some authors (Hitchcock et al., 1969; Reed, 
1954). Their combined range covers the western half of the United 
States. Gupta (1957) reported the latter two species to be distinct 
in epidermal pattern and stomatal frequency* however, we found 
all three to be essentially the same in these characters- 

In all species the cells toward the base of the pinnae have 
smoother walls than do the more distal ones; the former grade 
into the square or rectana:ular cells of the rachis epidermis. 

Epidermal streaks 

We examined only aerial leaves and floating leaves, so we 
did not find the ecological differences in epidermal wall patterns 
of aerial versus submerged leaves reported by Gupta (1962) 
and Gaudet (1964a). The latter are said to have smoother cell 


American Fern Journal 

walls than the aerial leaves. We found essentially no differences 
in epidermal cell walls between floating and aerial leaves. Van 
Cotthem's drawings of M. quadrifolia (1970, fig. 535) show more 
elongate cells with more deeply wavy walls than in our specimens. 

Marsilea ufpkr epidermis. Fig. 7. M. tenxjifolia. Fig. 8. M. fournieri 
Fig. 9. M. vncinata. Fig. 10. M. macropoda. Fig. 11. M. mollis. Fig. 12 


The presence of stomata on both surfaces of his specimen suggests 
it was not floating and that perhaps in this case floating and 
aerial leaves display some^Yhat different epidermal patterns- 

One morphological feature which appears to be correlated 
with floating leaves is the presence of reddish streaks on the under 
surface of the leaflets. The presence of these streaks has been 

Leaf Epidermal Studies in Marsilea lOo 

used occasionally as a key character in determining .V. mexicana 
(Knobloch and Correll, 1962, p. 47). The streaks are also com- 
monly found in il/. quadnjolia and .V. polycarpa. They are not 
merely chance streaks of pigmentation, but have an anatomical 
basis. In these areas the epidermal cells have tliinner, smoother 
walls than do the surrounding epidermal cells (Fiy. 4). The pres- 
ence of the streaks is strictly correlated with the absence of 


stomata on the lower surface, suggesting that these are floating 
leaves. Leaves with stomata on the lower surface have no streaks: 

drifolia, M 


stomata and suggestions of streaks of thin-walled cells on the 
lower surface that can be seen under the microscope. 

The streaks are by no means constant for any of the species 
mentioned above, but they are mvich more common in these than 
in any others. They have been seen in four specimens of the 
ilf. oliyospora-vestita-mucronata complex from the western United 
States and in M. deflexa from South America. This feature fre- 
quently occurs in species commonly found in water, which natur- 
ally have floating leaves frequently. The other species generally 
live in intermittent pools or moist meadows that are inundated 
only infrequently, and so only rarely have floating leaves. 

The fact that the cells have thin, smooth walls suggests that 
because of their submerged condition these area.s are slow to 
differentiate and essentially remain in a juvenile condition. 
Whether these areas have a function is not known. 

Stomatal frequency 

Stomatal size and relative frequency were determined for 
the upper and lower surfaces of all species. Frequency was meas- 
ured in five samples from each surface by counting the stomata 

visible in an ocular reticle (area 0.0715 mm-). The averages are 

given in Table I. As noted above,, some plants have no stomata at 
all on the low^er surface. It would appear from this feature, from 
the appearance of the plants (long petioles, few or no hairs), 
and rarely from label data, that these leaves were floating on the 


American Fern Journal 

water. Gaudet (19G4b) found floating leaves of plants grown in 
culture to have no stomata on the lower epidermis. This is to be 
expected since the floating leaves of water lilies (Nymphaea) 
have stomata limited to the upper surface, and the submerged 
leaves of members of the Podostemaceae have no stomata on 
either surface. It is interesting to note, however, that totally sub- 
merged leaves of Marsilea will initiate stomata on the upper but 
not the lower surface (Gaudet, 1964a). 

Marsilea upper epidermis. Fig, 13. M. mucronata. Fig. 14. M, oligq- 

spoRA. Fig. 15. M. \ 


In spite of the location of the stomata on the upper surface 
and the frequent lack of stomata on the lower surface, the in- 
ternal anatomy of the leaves is of the traditional configuration, with 
a palisade layer under the upper epidermis and an open spongy 
mesophyll layer below. There are air spaces in the palisade layer 
directly beneath the stomata. 

Aenal leaves generally have an appreciable number of stomata 
on the lower surface, but the stomata are more abundant on the 
upper surface (see Table I). In only one specimen was the fre- 
quency greater below than above. Although the frequency of 
stomata on a single surface is fairly constant, it may vary from 
plant to plant in a species, and the relative number of stomata 
on the upper and lower surfaces may also vary. Presumably this 
mtraspecific variation is largely ecologically controlled. 

Leaf Epider.mal Studies in Marsilea 107 

Inter^specific variation in istomatal fretiucMuy is quite pronounced 
in some cases. The number of stomata per unit area (0.0715 mm') 
on the upper epidermis ranges from 13 to over 40, although 
most are below 30. 

Gupta (1957) noted that the species he examined could be 
distinguished in some cases on the basis of relative stomatal 
abundance. MarsiJea vestitaj for example, was reported to have 
three to four times as many stomata as many of the other species. 
Our data do not confirm this, for we did not find M. vestita to 
have an exceptionally large number of stomata. In fact, one 
specimen with larger than normal stomata had exceptionally few. 
Marsilea mexicana had more abundant stomata than any other 
species we examined. 

Table I. Stomatal Frequency and Size in Marsilea 

Frequency Size {Microns) 

Species cfc Collector Upper Lower Upper Lower 

M. BERTEROi A. Brauii — Puerto Rico 

Britton €t al. 8872 18. 2 17. 22. 5 26. 8 

SintenisSSOS 19.0 15.6 21.6 24.3 

M. FOURXiERi C. Chr.— Mexico 

Brandegee, 5 Sept 1893 30. 8 19. 20. 8 25. 

Brandegee, 16 Oct 1899 32. 4 25. 8 20. 2 23. 5 

Carter & Moran 5394 20. 8 18.0 21 . 1 24. 2 

PringleS434 17.8 10.8 27.9 29.6 

^yiggim6422 17.6 11.2 25.2 27.9 

M. macropoda Engelm. — Texas 

Palmer 11245 20. 2 16. 25. 2 25. 7 

Palmer 33856 12. 8 14.4 23. 2 26. 8 

M. MEXICANA A. Braun— Honduras & Mexico 

Glassjuan 2054 34. 2 1.8 22. 8 23. 5 

Pr ingle 2007:. 37.8 20.8 — 

M. MOLLIS Robins. & Fern.— Mexico 

Hartman604 22.8 ,12.4 21.6 - 24.1 

Palmer 13 22.2 14.0 ' 27.3 29.9 

_ m.;mucronata a. Braun—Kansas & Texas • 

it Harvi^ 1581 29. 4 17. 8 24. 9 29. 2 


Tharp, 30 May 1937 22. 2 17. 2 22. 7 26. 1 

108 American Fern Journal 

Table I. Stomatal Frequency and Size in Marsilea (Continued) 

Frequency Size (Microns) 

Species & Collector Upper Lower Upper Lower 

M. nashii Underw. — Bahamas 
Nash & Taylor I4II 22. 4 14. 30. 2 34. 6 

M. OLiGOSPORA Goodd. — Idaho & Wyoming 
Nelson & Machride 1168 26. 2 18.2 21.9 28. 

Payson & Payson 2920 24. 4 21. 22. 6 26. 4 

Porter & Porter 9S99 19.4 15.4 23. 8 24. 9 

M. POLYCARPA Ilook. & Grev. — Puerto Rico 
Briilon et al 6726 26. 2 17. 8 19. 7 20. 9 

M. QUADRiFOLiA L. — Ontario, Mass. & Cult. 
Miller 647 29.4 22.1 . — 

Moldenke 1^608 . 17. 8 6.8 22. 4 23. 6 

Seymour, Aug 1900 19.2 17.2 21.0 21.9 

M. TENUIFOLIA Engleiil. — Texas 
r/iarp, 11 Aug 1941 19.6 17.6 26.5 29.1 

M. UNCINATA A. Braun — Texas 
TAarp, 3 Nov 1928 23.0 19.2 20.7 . 26.5 

M. VESTiTA Hook. & Grev.— Idaho, Calif. & Utah 
Christ 19130 14.2 9.6 26.8 28.8 

Heller 11633 20.0 18.2 25.1 26.4 

TilldtlSS 19.8 16.0 33.5 38.2 

Stomatal size 

Stomatal sizes were determined by measuring 20 stomata on 
each surface. Averages are shown in Table I. In every plant the 
stomata of the lower surface were found to be larger than those 
of the upper epidermis. Generally the difference is only 1-3 n, 


specimen of M 

outstanding in 

their very small stomata {M . polycarpa) or very large stomata 
(M. nashii), but most species have stomata 23-30 n long. 

Within species there is generally relative uniformity, although 
considerable variation is seen in M. fournieri, M. mollis, and 
M. vesiita. A polyploid series has been reported in M. minuta of 
India (Mehra and Loyal, 1959), a species for which Gupta (1962) 
reports stomatal size differences. 

Leaf Epidermal Studies in Marsilea 109 




Our study has shown that considerable variation exists in the 
epidermis of Marsilea speoieSj at least some of which has taxonomic 
significance. However, in order to determine more definitely 
the meaning of this variation, plants need to be grown under 


controlled conditions to determine the effects of water and light 
on the numbers and location of stomata; more extensive sampling 
is needed to establish the limits of variation within species; and 
more cytological information would be helpful in assessing the 
role that polyploidy plays in controlling stomatal size. 

Literature Cited 

Gaudet, J. J. 1964a. Morphology of Marsilea vestita I. Ontogeny and morph- 
ology of the submerged and land, forms of the juvenile leaves. Amer. 
J. Bot. 51:495-502. 

•■ . 1964b. Morphology of Marsilea vestita II. Morphology of adult 

and submerged leaves. Amer. J. Bot. 51 : 591-597. 
Gupta, K. M. 1957. Some American species of Marsilea with special reference 

to their epidermal and soral characters. Madrono 14: 113-127. 
— . 1962. Marsilea. Botanical Monograph No. 2. Council of Scientific 

and Industrial Research, New Delhi. 
Hitchcock, C. L., A. Cronqtist, M. Ownbey and J. W. Thompson. 1969. 

Vascular Plants of the Pacific Northwest, Ft. 1. Univ. of Washington 

Press, Seattle. 

Knohloch, I. W. and D. S. Correll. 1962. Ferns and Fern AUies of Chihua- 
hua, Mexico. Texas Research Foundation, Renner. 

Mehra, p. N. and D. S. Loyal. 1959. Cytological studies in Marsilea with 

particular reference to Marsilea minuta L. Res. Bull. Panjab Univ. 
n.s., 10: 357-374. 

Reed, C. F. 1954. Index Marsileata et Salviniata. Bol. Soc. Brot. II, 28: 1-61. 

Van Cotthem, W. 1970. Comparative morphological study of the stomata in 

the Filicopsida. Bull. Jard. Bot. Nat. Belg. 40: 81-239. 

The New York Botanical Garden, Bronx, NY 10458. 

IIU American Fern Journal 

The American Species of Plagiogyria sect. Carinatae^ 

David B. Lellinger^ 

Plagiogyria is principally a genus of China and adjacent regions, 
and is usually said to have about a dozen species in the New World. 
These species and their nomenclature have never been adequately 
studied in the past, although Copeland (1929) attempted to 
distinguish them and named several new species. Ching (1958) 
has written an extensive treatment of the species found on the 
Asiatic mainland. 

Among ferns, Plagiogyria is unusual in having few or no tri- 
chomes at maturity. Young portions of the plants are covered 
with mucilage which is secreted by many, deciduous or incon- 
spicuous, glandular hairs. In the Asiatic species studied by Hennip- 
man (1968), the hairs are usually less than 1 mm long. Scales are 
absent, although patches of matted hairs and mucilage sometimes 
resemble the "amorphous" scales characteristic of a few species 
of Blechnum. Plagiogyria is unique among the leptosporangiate 
ferns in having sporangia with complete, oblique annuli and about 
48 spores in each sporangium. The characters of the gametophyte 
are predominantly primitive, according to Stokey and Atkinson 
(1956). It is clearly an isolated genus, and in recent years has been 
put in its own family or order. 

The sterile fronds of the American species of sect. Carinatae, 
which supply all of the key characters, vary considerably within 
each species. The differences between the species are subtle, and 
immature or aberrant specimens may be difficult to place accu- 
rately; there seem to be no useful characters beyond those men- 
tioned m the keys. Because of the size overlap between extreme 
specnmens of the various species and the general similarities of 
their fronds, descriptions are of little use, and so have not been 
included in this paper. 

^ Partial cost of publication of this paper has been borne by the Smithsonian 

* I wish to thank the curators of the herbaria from which T borrowed or 
examined s{)ecimens for their cooperation. 

American Species of Plagiogyria sect. Carinatae 111 

Most of the New World species of Plagiogyria are quite re- 
stricted in distribution: one is from Hispaniola, one from Jamaica, 
one from Mexico^ one from Costa Rica to Bolivia, and one from 
southeastern Brazil. The only widespread American species, P. 
semicordata i is found in Cuba and Jamaica and from ^Mexico 
southward to Venezuela and Bolivia. All of the species are ter- 
restrial at middle and upper altitudes in moist, cool mountains 
and are absent from coastal and interior lowlands. 

Plagiogyria sect. Carinatae Ching ex Lellinger, sect, nov.^ 

Terrestrial ferns up to ca. 1 m tall, with erect, dimorphic fronds 
clustered at the tip of erect, woody, atrichomatous, dictyostelic 
rhizomes up to ca. 15 cm long and 1.5 cm in diam. (exclusive of the 
crowded and persistent stipe bases). Stipes and rhachises stramin- 
eous, brownish near the stipe base. Stipes 5-30 cm long, usually 
triangular, sulcate, only slightly expanded at the base; vascular 
bundles 3 at the base, fusing into a single V-shaped and then an 
U-shaped trace, this becoming nearly ribbon-shaped in the rhachis; 
aerophores obsolete on the dorsal surface of the stipe bases or 
absent. Fronds devoid of scales, with mucilage-producing, glandu- 
lar hairs when young, deeply pinnatifid (or the fertile sometimes 
pinnate), usually lanceolate, linear-lanceolate, or narrowly lan- 
ceate; rhachises terete below, sulcate above. Sterile fronds her- 
baceous or subcoriaceous; pinnae up to 15 cm long, (3)5-10(12) 

* Until now the name Plagiogyria sect. Carinctoe and the names of the sub- 
sections proposed by Ching (Acta Phytotax. Sinica 7: 135. 1958) have been 
invalid. Although they were in proper form and were provided with Latin 
diagnoses, the conditions of Art. 37 of the Code, which became effective about 
five months before Ching published, were not fulfilled- I therefore wish to 



Lomaria adnata Blume [= Plagiogyria adnda (Blume) Bedd.]. 

Plagiogyria subsect. Euphlebiae Ching ex Lellinger, subsect. nov. Type 
species: Lomaria euphlebm Kunze [ =^ Plagiogyria eu] hlebia (Kunze) Mett.]. 

Plagiogyria subsect. Pycnophyllae Ching ex Lelhnger, sub£e?t. nov. Type 
SPECIE'S: Lomaria pycnophylla Kunze [== Plagiogyria pycnophylla (Kunze) 


Ching also proposed sect. Eaplagiogyraej which is superfluous; according to 
Art. 22 of the Code, the typical section must be an automatic tautonym 
repeating without citation of authority the name of the genus, i.e., sect. 


A..MERICAN Fern Journal 

mm wide^ the margins subentire or crenate to sharply serrate or 
doubly sOj the veins solitary or paired at the base and simple or 
1-forked (see Fiys. 1-5). Fertile fronds herbaceous, generally 
longer and narrower and with longer stipes and shorter laminae 
than the sterile fronds; piiuiae up to 12(15) cm long, 2-4 mm wide, 

Diagrams of Venation types in the American sPEriES of Plagiogyria 
SECT. Carinatae. Fig. 1. Simple. Fio. 2. Paired at the base. Fig. 3. Forked 



the margins entire or irregular, often thin, somewhat involute, the 
veins 1-forked and sometimes uniting with an adjacent vein at the 
tip or forming a continuous commissure; receptacles somewhat 
enlarged and thickened; sori elongate; sporangia large, the stalk 
long, of 4-6 rows of cells, the capsule laterally compressed, the 
annulus obliciue, continuous; spores 48 per spoiangium, tetra- 
hedral, tuberculate to nearly smooth; x = ca. 25. 

Type species: Plagiogyria aryutissima Christ. 

American Species of Plagiogyria sect. Cartnatae 113 

Key to the American Species of Plagiogyria sect. Carixatae 

Pinnae of sterile fronds 5-7(8) mm wide; veins simple and either solitary or 

paired at the costae, or veins simple and 1-forked (Figs. 1-3). 
Margins of sterile pinnae simply aerrate, the veins beneath usually at 

least slightly prominulous (Mexico) ; 2. P, pedinala 

Margins of sterile pinnae crenate to doubly serrate, the veins beneath 

not prominulous (Brazil) 3. P. fialhoi 

Pinnae of sterile fronds (7)8-12 mm wide; veins solitary and l-forked or 

paired at the costae and 0-2-forked {Figs, 2-5). 
Sterile fronds decidedly narrowed at the base, the basal pinnae no more 

than half as long as the longest pinnae. Sterile fronds 6-14(22) 
cm wide,^ the pinna margins usually biserrate, at least toward the 
apex (Jamaica, Cuba, Mexico to Venezuela and Bolivia). 

\. P. semicordaia 

Sterile fronds not much narrowed at the base, the basal pinnae about 

two-thirds as long as the longest pinnae. Sterile fronds 10-24(S0) 

cm wide. 
Sterile fronds (10)13-24(30) cm wide^ (Costa Rica to Bolivia). 

4, P. cosiaricensis 

Sterile fronds 10-15(18) cm wide. 

Sterile pinnae crenate-serrate mostly toward the apex, sub- 
entire toward the base; laminae mostly 15-18 cm wide 
(Jamaica) 5. P. maxonii 

Sterile pinnae sharply and evenly serrate throughout; laminae 

mostly 10-15 cm w^ide (Hispamola) 6. P. serndafa 

1. Plagiogyria semicordata (Presl) Christ, Farnkr. 176. 1897. 

LomaridhimsernkordnUtm Presl, Epim. Bot. 155. 1849 [1851]. Type: Colom- 
bia, sin coll. (presumably PR or PRC not seen). 

Plagiogyria biserrata Mett. Abhandl. Senckenb. Naturforsch. Gesell., 
Frankfurt 2 [Farng. Flag. 8]: 272, t. 15, f. 1-18. 1858. Lbctotype: "Reg. 
alpina/' Colonia Tovar, Venezuela, Moriiz 4OO (B not seen Morton photo 
10211; isolectotypes HBG not seen Morton photo 5339, US), chosen by 
Copeland (1929, p. 409). The other syntypes are Mexico, Karwinsky (B 
not seen) and ^^Hatites Andes de Truxillo et de Merida," alt 4000-4500 
ft, Linden 556 (B not seen; isosyntype BR Weatherby photo US)AThe 
latter specimen I identify as P. costaricensis. 

* Sterile fronds &-8(12) cm wide in South America, 8-12(15j cm wide in the 
West Indies, and 8-14 cm wide in Central America, with some specimens from 
Oaxaca and Morelos, Mexico, up to 22 cm wide. 

' Sterne fronds 12-20 cm wide in South America, (9)12-15(30) cm wide in 

Costa Rica. 

114 American Fern Journal 



Plagtogyria obtum Copel. PhU. J. Sci. 38: 413. /. 13,1929. Type: Near Pico 
Turquino, Cuba, Leo.t 11126 (US; isotype NY|. 

Range and h\bitat: Mountains of Cuba and Jamaica (alt 1650- 
2220 m), of Mexico (Morelos and Puebia to Chiapas), Guatemalaj 
Ccs'a Rica, and Panama (alt 2500-3150 m), and of Venezuela, 
Colombia, Ecuador, Peru, and Bolivia (alt 2500-3500 m), in 
moist, often shady places in coniferous or hardwood forests and 
subparamo areas. 

Pla^fiogyria semicjrdata has biserrate pinnae that are narrower 

than these of P. costaricensiSy which are broader and denticulate. 

Ihe sterile pinnae of P. pectmata, in contrast, are simply serrate. 

The apex of the sterile fronds of P. semicordata is more tapering 

than is that of P. costaricensis. It is possible that most of the 

South American specimens I have identified as P. semicordata 

particularly those not from Venezuela — are small, contracted, and 

somewhat aberrant specimens of P, costaricensis, but I have no 
proof of this. 

Representative specimens examined: Cuba : Ortente: Pico Turquino, alt 
2000 m, Seifriz 1001 (US), northern spur, alt 1850 m, Ekman 5465 (NY, US), 
alt 2040 m, Ekman 5496, I4546 (both US). Jamaica : Blue Mountain Peak, alt 
2100-2225 m, Maxon 1413 (US), Underwood 2538 (NY), summit, alt 2100-2200 
m, Maxon 9875 (NY, US); Between Portland Gap and Blue Mountain Peak, 

alt 5500-7400 ft, Wilson cfc Wehder 623 (US); unlocalized, Gilbert 225 (NY), 
Hart 188 {V^). . " . . 

I q 

Mexico: Veracruz: Orizaba, MiiUer in 1855 (NY). Puerla: Honey Station, 
Pringle 15703 (F, US). Morelos: Zempoala, Lyonnet I4I9 (US). Oaxaca: 




HuEHUETENANGO: Cerro Canana, between Nucapuxlac and Canan.4, vSierra de 
los Cuchumatanes, alt 2500-2800 m, Sleyermark 49090 (F, US). Costa Rica: 
Cartago: S slope of Volcdn Turrlalba, near the Finca del Volcdn Turrialba, 
alt ca. 2000-2400 m, Standley 35133 (US) ; Behind the Hotel La Georgina, near 
Vaia MUls, alt 2900 m, de la Sola 5100 (US), alt 3000 m, A. Jimmez M. S^Ol 
(F). San Jos^: Cerro de las Vueltas, alt 3000 m, Standley 43883 (US) . Panama : 
Volcan Chiriqui, above El Potrero Camp, alt 2890-3025 m, Maxon 6334 (US). 

Venezuela: Aragua: Near Colonia Tovar, Fendler 325 (F, NY). Colombia: 
Antioquia: Highland of Santa Rosa, alt 2500-2700 m, Lehmann (US). Cundi- 
namarca: Macizo de Bogotd, Quebrada de Chico, alt 2800-3000 m. Cuatre- 

American Species of Plagiogyria sect. Cartxatae 115 

cams 5511 (US); Near La Gruta, Bogota, Litih & Uttle 7848 (US); Gtmdalupe, 
5 km E of Bogota, Liltle <fc Mile 9184 (US), Ho ught 5659 (F, NY, US). Ecuador : 
Azuay: Paramo de Tinajillas, 30-50 km S of Cuenca, alt 11000-11500 ft, 
Camp E-2225 (F, NY, US). Peru: Amazon as: Summit of Puma-urcu SE of 
Chachapoyas, alt 3100-3200 m, Wurdack 1159 (F, NY, US). Bolivia: La Paz: 
North Yungas, Unduavi, alt 3500 m, Buchtien 2I46 (US). 

2. Plagiogyria pectinata (Liebm.) Lellinger, comb, nov, 

Lomaria pectinata Liebm. Vid. Selsk. V, 1 : 233 [seors. 81]- 1849. Type: Cerro 
de Sempoaltepec, Oaxaca, Mexico, alt 8000-9000 ft, Liebmann (C not 
seen) . 

Lomaria arguta Fee, Mem. Soc. Hist. Nat. Strasb. 5 [8 Mem.]: 70. 1857. 
Type: 'Tallee d'Orizaba,'* Cerro del Agua, Veracruz, ^Mexico, alt 2700m, 
Schaffner 98 (P? not seen). 

Plagiogyria aequidentaia Fourn. Mex. PI. 1: 133. 1872. Type: Orizaba, 
Veracruz, Mexico, F. Miiller 723 (Hb. Lenormand not seen). The brief 
description, which contrasts the simply serrate pinnae with P. biserrala 
Mett. (a synonym of P, semicordata) ^ is sufficient to place the name. 

Plagiogyria arguia (Fee) Copel. Phil. J. Sci. 38: 407. 1929. 

Range and habitat: Hillsides and mountains of northern and 
central Mexico (Chihuahua to Jalisco, Alorelos, and Puebla), 
at 1750-3200 m altitude, in shaded, moist places in coniferous and 

coniferous-hardwood forests. 

Generally the basal pinnae of the sterile fronds of P. pectinata 
are about half as long as the longest pinnae, but in specimens 
from Oaxaca, lilexico, the sterile fronds have the lower pinnae 
even more reduced. 

Additional specimens examined®: Mexico: Chihuahua: Near La Rocha, 


NE slope of Sierra Mohinora, alt 7500 ft, Correll & Gentry 23112 (US); AIo- 
jarachic, Knobloch 594? (US). Durango: San Ramon, Palmer 124 (NY, US). 
Jalisco: E of Mamantlan ca 15 mi SSE of Autldn by way of Chante, alt 8300 
ft, WUbur * WUhtir 2001 (US); near Etzatlan, Rose & Painter 7598 (NY, US). 
Hidalgo: Beside the Rio Teponapa, vicinity of Zacualtipan, alt 2000 m, 
Moore 3207 (US). Puebla: Near Honey Station, alt 5800 ft, Pringle 8958 
(F, NY, US). MoRELOs: Lagunas de Zempoala, Bro. Sanchez S. 466 (US). 
Mexico: La Juntas, Sultepec Road, alt 3200 m, Lyonnet3119 (US); Zempoala, 
alt 3200 m, Maluda 281 40 (US) ; Cerro de Acatitlan, alt 2000 m, Matuda 30762 

• The specimens (from Oaxaca south to Chiapas and adjacent Guatemala) 
differ from typical P. pectinata in having the lower pinnae of their sterile 
lanfiinae reduced gradually below to 0.5-1 cm long, or even to mere auricles; 
typical specimens (from Oaxaca north to Chihuahua) have the lowest pinnae 

0.35-0.65 as long as the longest ones. 


American Fern Journal 

(US) ; Crucero, 11 ^ovl9U, Hinton 6823 (NY, US); Los Hornos, alt, 2700 m, 



llinas, alt 9000 ft, C. L. Smith 2063 (NY, US) ; Valley of the Rio Yelagago, ca. 
20 mi. NE of Villa Alta, alt 6800-7200 ft, Mickel 929 (US) ; N slope of Cerro 
Felon, along road from Ixtlan toward Tuxtepec, alt 9000 ft, Mickel 11^2 (US). 
Chiapas: Zonehuitz, Munch 102 (US); between San Cristobal las Casas and 



mala: Huehuetenango: Top of Cerro Chemalito, Sierra de los Cuchuma- 
tanes, 3.5 mi. W of Santa Eulalia, alt 3100-3150 m, Sleyermark 49927 (F, US). 

3. Plagiogyria fialhoi (Fde & Glaziou in Fee) Copeland, Univ. 

Calif. PubL Eot. 19: 297. 1941. 

Lomnria fialhoi Fee & Glaziou in Fee, Crypt. Vase. Bres. I: 239, /. 7. /. 2, 
1869. Type: Serra os Orgaos, Est. Rio de Janeiro, Brazil, Glaziou 3326 
(P not seen). 


Distribution and habitat: Southeastern Brazil, in the high 
portions of mountains, in low forests, according to £ehnem (1967). 

Additional spec mens examined: Brazil: Santa Catarina: Source of the 
Rio de Oratorio, alt 1200 m.^Smith & Reitz 10232 (US); Campo dos Padres 
between Fazenda Santo Antonio and Fazenda Campo dos Padres, alt 1400- 
1650 m, Smith it lieitz 10377 (US). 

4. Plagiogyria costaricensis :Mett. ex Kuhn, Linnaea 36: 149. 


Plagiogyria anit>odonta Copel. Phil. J. Sci. 38: 409, /. 9. 1929. Type: Dead 
crater of Volcdn Poas, Prov. Alajuela, Costa Rica, alt 2800 m, 0. Jimenez 
1018 (US). Par.\types: Volcan Poas, Prov. Alajuela, Costa Rica, upper 
slopes between the Hotel and the crater, alt 2500-2640 m, Standley 34S95 
(US, , summit, alt 2644 m, Tondiiz 10714 (US). 

Plagiogyria denticulcta Copel. Phil. J. Sci. 38: 412, /. 12, 1929. Type: 'Im 
Nebelwald des Bergkammes der Laguna verde bei Comparapa [Com- 
arapa]," Depto. Santa Cruz, Bolivia, alt 2600 m, Herzog 1954 (US). 
Paratype: North Yungas, Bolivia, alt 3300 m, Btichlien 1908 (US). 

Plagiogyria latifolia Copel. Phil. J. Sci. 38: 411, L IL 1929. Type: Cani, 7 
mi. NE of Mito, Depto. Hudnuco, Peru, alt ca. 8500 ft, Macbride 3432 


Type: Volcan Barba, Prov. Heredia, Costa Rica, Wendland 1066 

(C? not seen) A drawing of the holotype (B) illustrates the broad, 

coarse, sterile lamina and the fertile lamina only slightly reduced 
at the base. 

American Species of Plagiogyrta sect. Carinatae 117 

Distribution and habitat: ^Mountains of Costa Rica and the 
Andes, above (1200)1800 m altitudej in wet woods and shrubby 

This species is larger and coarser than P. semicordata. The 
costae beneath often have what look Hke rather amorphous scales 
along them, but these are merely dried up patches of mucilage 
which have trapped numerous deciduous hairs. Sometimes a few 
cells from deciduous glands can also be seen; their terminal cells 
are swollen and reddish-brown. 

Additional specimens EiXamined: Costa Rica: Alajvbla: Volcan Poas, 
near the summit at the edge of the crater, Burger 4352 (NY), upper lagoon^ 
alt 2564 m, Piitier 2970 (US), Poas Lake, alt 8590 ft, Etork 23S8 (US , along 
the road to the volcano, alt 6000-7000 ft, Scanwian 7607 (US). Heredia: 
Volcdn Barba, alt 2700 m, Valerio I46 (US). Cartago: La Pastora, near 
Volcdn Irazu, alt 9000 ft, Stork 2378 (US,; N of Volciin Irazu, alt 10000 ft, 
Stork 1284 (NY); Volcdn Turrialba, alt 3100 m, Torres R. 12 (US'. S\N Josk: 
Cerro de las Vueltas, alt 2700-3000 m, Stmdley & Valeria 43S40 (US), alt 
3000 m, Pittier 10498 (US, ; Lagima de la Chonta, NE of Santa Maria de Dota, 
alt 2000-2100 m, Standleij 42129 (US). 

Venezuela: M^Irida: Upper Andes of Truxillo and M^rida, Culata, alt 1600 
toises [= 10240 ft], Linden 666 [a syntype of P, hiserrcta Mett.] (B not seen; 
BR Weatherby photo USj. C olcmbia : Cundinamakca: SE of Bogota, alt 2800 
m, Andre 976 (NYj; Usaquen, near Bogotii, alt 2700 in, Uribe U. 208 (F); 
Quebrada de las Delicias, Macizo de Bogota, alt 2650-2770 m, Cuatrecasaa 
6466 (F, US) ; Summit of Monserrate, E of Bogota, LMle & Liiile 81S9 (US) ; 
Western slopes of Paramo de Cruz Verde, alt 3150 m, Cualrecasas 402 (US). 
Narino: Headwaters of the Rio Tellez, above Soledad, alt 2900-3150 m, 
Ewan 16513 (US). Peru: HuiNUCOt Yunano, alt ca 6000 ft, Mathride 3830 
(F, US). Bolivia: La Paz: North Yungas, Unduavi, alt 3200 m, Buchlien 
410 (F, NY), alt 3300 m, Buchlien 2166 (US), 2677 (F). 

5. Plagiogyria maxonii Copel. Phil. J. £ci. 38: 411, L 11. 1929. 

T\pe: Summit of Blue Mountain Peak, Jamaica, alt 21CO-2200 
m, Maxon 9910 (US; isotypes F, NY). 

Distribution and habitat: Mountains of Jamaica, in wet 
forest above 1700 ni altitude. 

Additional specimens examined: Jamaica: Sir Johns Peak, Watt 6 in 1910 
(NY, US); Mossman's Peak, upper eastern ridge and summit, alt 1700-1925 
Hi, Maxon 9721 (F, NY, US); Blue Mountain Peak, alt 6500-7325 ft, Under- 
wood 1495 (NY), alt 7000-7423 ft, Maxon 1439 (US), Underwood 2654 (NY), 
alt 7420 ft, Harris 7144 (F). 

118 A:\rERicvN Fern Journal 

6. Plagiogyria senulata (Willd.) Lellinger, comb. nov. 

AcrosUchum serrulaium Willd. Sp. PL ed. 4, 5: 113. 1810. Based on Plate 81 y 
Polypodium fuscuiii tenuisshnis deniiculis serratum, of Plumier's "Tracta- 
tus de Filicibus Americunis." Plumier's plate illustrates several char- 
acteristics of this species (or of Plagiogyria in general): the evenly serrate 
pinnae with 1-forked veins, the acrostichoid fertile frond, and the sterile 
fronds scarcely narrowed below. His description mentions the blackish 
rhizome and the rhizome hairs, which are also characteristic of Plagio- 
gyria, but not of Polypodium^ to which Plumier ascribed his plant. He 
gives the locality as "la bande du Sud au Grand Cul-de-sac/ proche un 
endroit que les Boucaniers appellent le fond ^pineux" [Haiti]. 

Blechnum urbanii Brause in Urban, Symb. Antill. 7 : 157. 191 1, as "urbani.^^ 
Type: Near Constanza, Santo Domingo, alt 2200 m, von Tiirckheini S4O8 
(B not seen photo US, fragment US). 

Plagiogyria urbanii (Brause) Copel. Phil. J. Sci. 38: 413. 1929. 

Distribution and habitat: ^Mountains of Hispaniola, in 
mossy and marshy places in pine forests above 1850 m altitude. 

Additional specimens examined: Dominican Republic: Sierra de Ocoa, 
San Jose de Ocoa, Loma Sucia, alt ca 2300 m, Ekumn H12002 (US) ; Cordillera 
Central, Moncion, high ridge between the Rio Cerrobi and the Rio San Juan, 
alt ca 1900 m, Ekman HI 2790 (US); Sabana Nueva, between Piedra del 
Aguacate and Rio del Oro, alt 6200 ft, Howard & Howard 9123 (NY, US). 
Haiti: Massif de la Hotte, Torbec, top of Morne Formon, Ekman H7501 
(NY, US). 

Literature Cited 

Ching, R. C. 1958. The fern genus Plagiogyria on the mainland of Asia. Acta 

Phytotax. Sinica 7: 105-154, /. xxviii~xl 

t 1-15. 



some leptosporangiate ferns. Bhmiea 16: 97-103. 
Sehnem, a. 1967. Plagiogiriaceas in R. Reitz, Flora Ihistrada Catarinense, 

Parte I. Itajai, Brazil. 
Stoket, Alma G. and Lenette R. Atkinson. 1956. The gametophytea of 

Plagiogyria glauca (Bl.) Mett. and P. semicordata (Pr.) Christ. 

Phytomorphology 6: 239-249. 

U. S. National Herbarium, Smithsonian Institution, 
Washington, D. C. 20560. 

The Genus Stenochl.\ena 119 

The Genus Stenochlaena J. Smith 
with Description of a New Species 


The genus Stenochlaena in Chiistensen's "Index Filicum"(1905) 
included species belonging to three distinct genera. The three 
are superficially alike in having simply pinnate fronds with 
entire pinnae and free veins, the fertile fronds having narrow 
pinnae almost completely covered beneath with sporangia. But 
if one examines the vascular anatomy, scales and spores, one 
finds the superficial similarity hides a great diversity. The three 
genera are Stenochlaena s.str., Lomario-psis Fee, and Teratophylhim 
Mett. I studied these plants in the forests of Malaya, and wrote a 
paper describing their differences (Holttum, 1932). The differ- 
ences are also summarized by Copeland (1947), and the species of 
the three genera in the INIalay Peninsula are described in Holttum, 
1954. The genera Stenochlaena and Teratcphyllum are confined 
to the Old World, but Lomariopsis is pantropic. Ihus all tropical 
American species formerly included in Stenochlaena belong to 
Lomariopsis (see Holttum, 1939), but Vareschi (1968) still places 
them in Stenochlaena, and his generic description includes a 
statement about vascular anatomy which applies to Stenochlaena 
s.str. and is not true of the species he describes. In 1932 I expressed 
an opinion that Stenochlaena is related to Acrcstichum (both 
genera have an exceedingly complex vascular anatomy) and thus 
to Pteris, and I still think this possible; but Copeland has included 
Stenochlaena in the Blechnaceae. Lcmaricpsis and Terattphyllum 
appear to me to be related to Tectaria and Cienitis. 

Six species of Stenochlaena are here recognized, two in Africa, 
the others in Asia, iVIalesia, and the Pacific. One of the latter 
has not yet received a valid name; as pointed out by Morton 
(1970) my attempt to give it one ignored the terms of the Inter- 
national Code of Botanical Nomenclature. In my paper of 1969 I 
also reported that examination of a type specimen in Presl's 
herbarium necessitated a re-interpretation of his name SJuJandi- 

120 American Fern Journal 

folia and therefore a re-naming of specimens previously so called 
by me. As a comprehensive account of all species of the genus as 
now understood has never been published, I take this opportunity 
of providing one^.to include the information of my paper of 1969. 
Only important synonyms and brief species descriptions are in- 
cluded. Further details may be found in the other literature cited. 

Key to the Species of Stexochlaena 

Fertile fronds bipinnate, the sterile sini])ly pinnate 1. S. tennifoha 

Fertile and sterile fronds both simply pinnate. 

Pinnae conunonly 15 pairs or more, the sterile pinnae to at least 15 X 3 

cm; fertile pinnae with very narrow sterile margin. 
Scales on tip of rhizome narrow, spreading; bases of pinnae not 

articulate . . .' 2. S. mildhraedii 

Scales on tip of rhizome almost circular, appressed; pinnae articulate 

or not. 
Lower pinnae at least, appearing fully articulate when dry; 

pinna-bases cuneate; fertile pinnae 2-3 mm wide; 

spores evenly tuberculate throughout. . .3. S. palustris 

Pinnae not distinctly articulate; sterile pinnae with broadly 

rounded to cordate bases; fertile to at least 5 mm wide; 

spores with raised dt broken ridges. 
Sterile pinnae broadly rounded at base, sometimes partly 

articulate; fertile pinnae 5 mm wide; spores with 

irregular broken ridges 4. S. milnei 

Sterile pinnae cordate at base, not at all articulate; fertile 

pinnae 7-10 mm wide; spores with parallel con- 
tinuous or broken ridges 5. S. cumingii 

Pinnae to 7 pairs; sterile pinnae to 10 X 2.2 cm; fertile pinnae 3 mm wide 

with thin reflexed sterile margins 1 mm wide 6. S. areolaris 

1. Stenochlaena tenuifolia (Desv.) ]Moore, Gard. Chron. 

1856: 193; INIorton, Amer. Fern Journ. 55: 165. 1965; 
Tardieu in Humbert, Fl. Madag., Fam. 5, 1: 110. 195S. 

Loniaria lenuiJoHa Desv. Berlin Naturf. Freund. Mag. 5: 326. 1811. 

Rhizome to 12 mm diameter; scales on rhizome-tip narrow, 
spreading; sterile pinnae very variable in size, to 30 X 3 cm, not 
at all articulate to rachis; fertile pinnae to 15 cm long with 10 12 
pairs or more of pinnules ca. 5 cm X 2 mm. 

DiSTKiBUTiox: East tropical to South Africa, :Madagascar, 

The Genus Stenochlaexa 121 

2. Stenochlaena mildbraedii Brause, Bot. Jalirb. Engler 53: 

384. 1915; Tardieu in Flore du Cameroun 3: 353, pL 
XXXIV, figs. 5,6, 1964. 


Rhizome-scales elongate and spreading; stipe 100-150 cm; 
frond 100-150 cm long; sterile pinnae" 16-25 pairs, to 40 X 4 cm, 
not articulate to rachis; fertile pinnae 4-5 mm wide. 

Distribution: West Africa, Cameroons to Angola, eastwards to 
Uganda; climbing on Raphia palms. 

3. Stenochlaena palustris (Burm. fil.) Bedd. Ferns Brit. Ind. 

Siippl. 26. 1876. Handb. Ferns Brit. Ind. 421. 1883. 
Holttum, Rev. Fl. Malaya 2 : 412, fig. 241. 1954. 

Polyopdium palustre Burm. fil. Fl. Ind. 234. 1768. 
Lomaria ? jnglandi folia Presl, Rel. Haenk. 52. 1825. 
Stenochlaena fraxini foil a Presl, Epim. Bot. 163. 1851. 


S, laurifoUa Presl, ihicL 164 (see Holttum, 1969, pp. 14, 51). 

Rhizome long-creeping or climbing, ca. 5-7 mm diameter; 
stipes 15-20 cm long containing up to 40 vascular strands; fronds 
40-80 cm long with 8-15 pairs pinnae; sterile pinnae articulate to 
rachis, very variable in sliape and size, commonly ca. 15 X 3 cm, 


exceptionally to 24 cm long but not proportionately wider, the 
base narroAvly to broadly cuneate;. fertile pinnae commonly 20 
cm long and 2-3 mm wide, with a very narrow, not reflexed, 
sterile margin; spores evenly tuborculate throughout. 

Distribution: India through Malesia to N. E. Australia; in 
the Pacific to Samoa and Tonga. 

4. Stenochlaena milnei Underwood^ Bull. Torrey Bot. Club 

33 : 38. 190G. 

S. juylandifolia sensv Holttum, Card. Bull. Str. Settl. 9: 139. 1937, non 
Presl; Copel. Feru Fl. Philip. 428. 1960. 

Kesembling *S. palustris but sterile pinnae always broadly 
rounded at base and articulation to rachis not complete; stipe 
of large fronds containing a larger number of vascular strands; 
fertile pinnae ca. 5 mm wide; spores bearing irregular broken 

Type: Solomon Islands, Milne 51S (K). 

DiSTRiBUTiox: Philippines, New^ Guinea, Solomon Islands. 

122 American Fern Journal 

When I was in New Guinea in 1963, I noticed that this was the 
common species of Stenochlaena in the neighborhood of Lae, not 
S. palustns. It may be that the fern mentioned by INIorton (1965, p. 
166) as cultivated in the University of Cahfornia Botanic Garden 
from spores collected in New Guinea is this species. Morton de- 
scribes it as a large and coarse vine; S. palustris has a more slender 
rhizome and smaller fronds, and I Avould not so describe it. 

5. Stenochlaena cumingii Holttum, sp. nov. 

S. laiirifoUa sensu Holtt. Card. Bull. Sir. Settl. 5: 259. 1932, non Presi; 
ibid, 9: 141. 1937; Copel. Fern Fl. PhUip. 428. 1960; Holttum, 1969, pp. 
51, 52. 

Stipes validus, basi fasces vasculares 100 vel ultra continens; 
pinnae steriles basi late cordatae, baud articulatae; pinnae fertiles 
7-10 mm latae; sporae earinis ± interruptis parallelis longitudi- 
nalibus ornatae. 


Type: Luzon, Cummg 226 (K; isotype BM) 

Distkibution: Moluccas, Philippines, New Guinea. 

This appears to me to be a very distinct species, though Copeland 
has expressed doubts. Stenochlaena milnei is almost exactly inter- 
mediate between >S. palustris and S. cumingii and may represent 
a hybrid. Cytological evidence is needed. 

6. Stenochlaena areolaris (Harr.) Copel. Philip. Joum. Sci. 


2C : 406. 1908; Fern Fl. Philip. 427. 1960. 

Rhizome slender; stipe of sterile frond ca. 15 cm, of fertile ca. 
30 cm long; sterile frond to 25 cm long with 7 pairs of pinnae; 
pinnae to 10 X 2.2 cm; fertile pinnae to 10 cm long, 3-6 mm wide, 
with thin margins 1 mm wide reflexed at right angles. 

Distribution: Philippines, New Guinea, climbing on Pandanus. 


Copeland, E. B. 1947. Genera Filicum. Waltham, Mass. 
HoLTTVM, R. E. 1932. On Stenochlaena, Lomariopsis and Teratophyllum in 

the Malay Peninsula. Gard. Bull. Str. Settl. 5: 245-323. 


— . 1954. A Revised Flora of Malaya, II Ferns of Malaya. Govern- 
ment Printer, Singapore. 

Spore Size and Germixatiox in Dryopteris 123 

HoLTTUM, R. E. 1969. A commentary on sorae type specimens of ferna in the 

herbarium of K. B. Presl. Nov. Bot. Inst. Bot. Univ. Carol. Prag. 

Morton, C. V. 1966. Observations on cultivated ferns, VIII. Stenochlaena. 

Amer. Fern. J. 55: 164-166. 

. 1970. Review of "A commentary on some type specimens of 

ferns in the herbarium of K. B. Presl.'' Amer. Fern J. 60 : 119-128. 
Vareschi, V. 1968. Flora de Venezuela, Vol. I. Instituto Botdn'co, Caracas. 


RovAL Botanic Gardens, Kew, Richafoxd, Surrey, Eng- 

The Variation in Spore Size and Germination 

in Dryopteris Taxa 

Dean P. Whittier and W. H. Wagner, Jr.^ 

Fern spores have provided a series of diagnostic characters for 
understanding the biology and relationships of the species. A 
number of studies have demonstrated, for example, that 32 spores 
in a sporangium are characteristic of apogamous species (Manton, 
1950). The size of spores within a genus is commonly related to 
the ploidal level of the species (Blasdell, 1963; Wagner, 1966; 
Whittier, 1970). Sterile hybrids tend to have a greater variation 
in spore size than normal sexual species (Wagner & Chen, 1965; 
Kanamori, 1969). It also has been demonstrated that some spores 
of sterile hybrids can germinate and produce gametophytes 
(Morzenti, 1967; deBenedictus, 1969), Recently, Weinberg and 
Voeller (1969) reviewed external factors that control germination 
of fern spores and, among other observations, ' they found that 
spores from the same plants taken at different times gave very 
different rates of germination. • 

^ Research supported in part by NSF Grant GB-8113. Mr. Johannes Jensen 
epared the spore materials for the experiments. 

124 American Fern Journal 


The wooclferns, Dryopteris, present one of the most difficult of 
American fern complexes because of their tendency to hybridize. 
A number of workers have recently studied these plants cytologi- 
cally and have shoAvn that many of the sexual taxa share one or 
more genomes with others. Except for D. fragrans, D, fiUx-mas, 
and D, marginalise all of the other sexual taxa have pairing affini- 
ties of the chromosomes with at least one other as shown in the 
following scheme (Wagner, Wagner, & Hagenah, 1969): 

D. "dilatata'' ' DD 

D. caynpyloptera DDII 

D. intermedia 


D. spinulosa IISS 

D, cristata SSLL 

Z>. ludoviciana LL 

D. celsa LLGG 

D. gcldiana - GG 

D. clintoniana SSLLGG 



Some of these, e.g., D. celsa, are evidently of hybrid origin, but 
they behave exactly hke normal species because of their balanced 
chromosome complements and normal spore and gametophyte 
production. Sterile hybrids in Dryopteris, on the other hand, 
have usually one or more ''unmatched'^ chromosome sets, and the 


resulting failure of these sets to pair normally produces spore 
abortion and sterility. 

The following investigation was undertaken to test the corre- 
lation of hybridity, spore size variation, and germinabiUty in 
several Dryopteris species and hybrids grown under unifonn 

Materials and Methods 

The Dryopteris spores wx^e obtained from plants in an experi- 
mental garden of the Matthaei Botanical Gardens at the Uni- 
versity of ^Michigan. Since the plants were grown under the same 
conditions, environmentally produced variations in the spores or 
germination would be reduced to a minimum. In addition, the 

Spore Size and Germination in Drvopteris 125 


cytology of the plants has been studied and the chromosome 
pairing at meiosis is known. The spores were collected July 22, 
1969, by the following method. A pinna was removed and was 
washed with water to remove the foreign spores. The washed 
pinna was placed in an envelope to dry and to release tlie spores. 

. A sample (40 spores) was obtained from each taxon and measure- 
ments were made with a calibrated ocular micrometer. The stand- 
ard deviation (Li, 1964) was taken as a measure of the variation 
in spore sizes. The total length of the spore, including the peri- 
spore, was measured because the endospore can not be observed 
in spores of some hybrids (Wagner &: Chen, 1965). The spores 
were sown on 2 ml of Knudson's solution of mineral salts at pH 
6.3 (Whittier, 1965) in capped, 1 dram vials. The cultural condi- 
tions included 12 hours of illumination (200 ft-c) per day and a 

temperature of 24 d= 1° C. Two hundred spores were observed 

seven days after sowing to determine the percentage of spore 

Results and Discussion 

Table I shows that the spores of the Dryopteris species had 
less variation in size, as measured by the standard deviation, than 
the spores of the Dryopteris hybrids. The germination of the spores 
of the Dryopteris species ranged from 78-95% with one exception. 
Spores of one plant of D. celsa had 37.3% germination. Hybrid 
spores had low percentages of germination with two exceptions, 
34.7% for D, celsa X marginalis and 36.0% for one plant of D. 
cliy\tcniana X cristata. These Dryopteris taxa demonstrate that 
a greater variation in spore size, which is mainly produced by 
spore abortion, is correlated with the sterile hybrids of the genus. 
This supports the conclusion reached by others (Wagner & Chen, 
1965; Kanamori, 1969). 

It is of interest to consider the taxa that had about 35%? germi- 
nation. This level of germination was between the high percentages 
for the fertile species and the low percentages for the hybrids. 
The reduced germination for the spores of one plant of D. celsa 
probably was due to an internal factor which influenced the spores 


American Ferx Journal 

or spore development. It is unlikely that the reduced germination 
can be explained by environmental conditions because these 
spores were formed, collected, and grown under the same condi- 
tions as spores from other plants of D, celsa. The high percentage 

Table I. Spore Size, Standard Deviation, and Percent Germination 


Tax a of 


Mean spore 
length in /* 









margin alis 

campyloptera X intermedia 

celsa X clintoniana 

celsa X goldiana 

celsa X marginalis 

clintoniana X cristata 
clintoniana X cristata 
cristata X intermedia 
cristata X marginalis 
cristata X spinulosa 
filix-mas X marginalis 
goldiana X marginalis 



± 4.2 

± 4.7 






• Germination 












D. celsa X marginalis and 

B. clintoniana X cristata was unexpected because low percentages 

germination had been reported 

& Chen 



tion for these two plants is unknown, but possibly it is due to a 


Spore Size and Germination in Dkyoptehis 127 


greater percentage of pseudomeiotic sporogenesis (Alorzenti, 
1962) in the sporangia of the plants. It should be noted that 
another hybrid, Z). cristata X marginaliSy also had a relatively 
high level of germination. Although in Dnjopleris the percentage 
of spore germination is generally large for the species and small 
for the hybrids, the exceptions make spore germination by itself 
a poor character for suggesting hybrid origin for individual plants. 



Literature Cited 

Blasdell, R. F. 1963. A monographic study of the fern genus Cystopt 

Mem. Torrey Bot. Club 21 (4) : 1-102. 
DeBbnedictus, V. M. M. 1969. Apomixis in ferns with special reference to 

sterile hybrids. Doctoral Dissertation, University of Michigan. 
Kanamori, K. 1969. Studies on the sterility and size variation of spores in 

some species of Japanese Dryopteris. J. Jap. Bot. 44: 207-217. 
Lr, J. C. R. 1964. Statistical inference. Vol. I. Edwards Bros., Ann Arbor, 

Manton, I. 1950. Problems of cytology and evolution in the Pteridophyta. 

Cambridge University Press, Cambridge. 
Morzenti, V. ]\I. 1962. A first report on pseudomeiotic sporogenesis, a type 

of spore reproduction by which "sterile'' ferns produce gametophytes. 

Amer. Fern J. 52 : 69-78. 
— . 1967. Asplenium plenum: a fern which suggests an unusual 

method of species formation. Amer. J. Bot. S4: 1061-1068. 
Wagner, "W. H., Jr. 1966. New data on North American oak ferns, Gymno- 

carpium. Rhodora 68: 121-138. 
— ^ and K. L. Chen. 1965. x\bortion of spores and sporangia as a tool 

in the detection of Dryopteris hybrids. Amer. Fern. J. 55; 9-29. 
, F. S. Wagner and D. J. Hagen.vh. 1969. The log fern (Dryopteris 

celsa) and its hybrids in Michigan — A preliminary report. Mich. Bot. 


Weinberg, E. S. and B. R. Voeller. 1969. E.\ternal factors inducing germi- 
nation of fern spores. Amer. Fern J. 59; 153-167. 
Whittier, D. p. 1965. Obligate apogamy in Cheilanthes tomentosa and C. 

alabamensis. Bot. Gaz. 126: 257-281. 

. 1970. The rate of gametophy tic maturation in sexual and apogam- 

ous species of ferns. Phy tomorphology 20 : 30-35. 

Department of General Biology, Vanderbilt University, 
Nashville, Tennessee 87203 and Department of Botany, 
University of Michigan, Ann Arbor, ^Iichigan 48104. 

128 American Fern Journal 

The Gametophytes of Natural Hybrids 

in the Fern Genus Pellaea 

Thomas R. Pray 

In the course of extensive field work relative to em investigation 
of Pellaea (Pray, 196Sa, b, 1970) a number of collections were 
made on the western slopes of the Sierra Nevada in California of 
an undescribed member of the genus. Analysis of the plant sug- 
gests that apparently it is a hybrid (which will be described in 
detail elsewhere) of P. hridgesii and P. mucronala. The various 
features of the plant are mostly intermediate between those of the 
morphologically dissimilar parental species. As is to be expected in 
such a Wide cross, the sporophyte is essentially sterile. At times, 
however, occasional individuals can be found to yield a few viable 
spores among the far more numerous nonfunctional products of 
abortive sporogenesis. Among the many individuals examined, 
two plants from a locality near Yosemite (Pray 1958, 1961) had 
sufficient functional sporangia to give a few spores to be sown on 
agar. In these sporangia, spore size was variable, with a few ob- 
viously larger than the rest. It was apparently the larger spores 
that were capable of germination. Accurate spore number per 
sporangium was not determined. It was, however, less than the 

normal 64. 

Spores were sown on nutrient agar as in previous investigations 
(Pray, 1968a). Most of the larger gametophytes were transferred 
to sterile soil 2 months after sowing to complete their development. 
The account which follows is necessarily incomplete due to the 
small number of gametophytes available. In order to determine 
whether the gametophytes could reach maturity and reproduce, 
most were allowed to continue growing, rather than be sacrificed 
for detailed microscopic examination. The usual complete develop- 
mental sequences were therefore not fully realized. Spores from the 
two hybrid plants used were sown separately. No differences were 
noted between the progenies, however; they are, therefore, de- 
scribed together. . 

Gametophytes of Natural Hybrids in Pellaea 129 


The outstanditig feature of the gamotophytes from the hybrid 
plants was their extreme variability. None of these variations can 
be attributed to crowding or other unfavorable growth conditions. 
The germinating spore produces a filament as in other Pellaeas 
(Pray, 1968a, b, 1970). The filament may then give rise to a regular 
spathulate plate phase (Figs, i, £) which lacks a true apical cell. 
A few thalli {Fig. 2) may show a terminal cell with the appearance 
of an apical initial. Such a cell does not function as a true initial, 
however. Growth is generally diffuse at this phase. As the plate 
expands a marginal meristem with multiple initials is differen- 
tiated. This may occur in a terminal position. If so, the meristem 
soon occupies a shallow sinus and a typical cordate thallus with 
more or less equal wings is the result {Fig, 12). Although this form 


of thallus, derived from the most regular of the young plates, was 
observed in both groups of gametophytes, it is found in 1% or 
less of the total number of thalli. Sometimes a somewhat lateral 
meristem will differentiate {Figs. 5y 7). Strictly lateral meristems 
were also observed {Fig. 10). It is assumed that these latter types 
will eventually give rise to cordate thalli through the formation 
of a second wing and unequal growth rates of the regions lateral 
to the meristem. Thus the meristem is shifted to a terminal 
position. Asymmetrical thalli such as Fig, 11 are usually the de- 
rivatives of those with at least sliglitly lateral meristems at an 
earlier stage. 

Except for those relatively few gametophytes which followed 
fairly standard pathways of development for Pellaea^ the re- 
mainder were f[uite aberrant. These usually lacked, or tended to 
lose, recognizable centers of growth. Frequently the shift from 


filamentous to two-dimensional growth failed to occur or was 
considerablj' delayed (Fiys. 3, 4). This inability to initiate orga- 
nized two-dimensional growth could be permanent with the result 
that the filamentous phase continued to spectacular lengths. 
Fi(jure 15 shows the major portion of a 2-month old gametophyte 
4 mm long. This individual has remained entirely uniseriate except 
for the vertical w^all near the base. Some of its more regular con- 

American Fern Journal 

Volume 61, Plate 18 

1 mm 


Figs. 1-12. Pellaea bridgesii X P. mucronata. Ftgs. 1-3, 5, 6, 9-11 

Pray 1861 (VSC). Figs. 4. 7. 8. 12 from Pray 

Gametophytes of Natural Hybrids in Pellae-\ 131 

temporaries were typical cordate forms up to 4-5 mm in width. 
Sometimes the filamentous phase did eventually evolve non- 
terminal 2-dimensional regions {Fig. 4) which could differentiate 
.a mofe normal appearing plate {Fig. 6). In this instance a meristem 
might finally appear although such was not actually observed. 


Figure 6 might be interpreted as a case in which the filamentous 
portion is a secondary outgrowth and not the original terminal 
portion of the thallus. Ihe precise developmental histories of ex- 
tremely peculiar, ameristic forms {Figs. 16, 17) is also uncertain. 
It is possible that these have arisen through localized expansion of 
median or basal areas of extended filaments. On the other hand, 
fairly regular early plate phases may have failed to differentiate 
meristems and developed filamentous outgrowths subsequently. 
It is worth noting that Figs. 4? 15-17, show much greater cell size 
than their counterparts. This suggests that continued cell enlarge- 
ment has occurred in the absence of normal rates of cell division. 
It is also possible, of course, that as a result of meiotic irregularities 
these particular thalli have higher chromosome numbers. 

Between the extremes represented by the above examples, there 
were intermediates of all sorts. Figure 5, with its large cell size, 
suggests a relationship to Fig. 4- Or perhaps this figure represents 
a form which has had more regular development and will ulti- 
mately organize a lateral meristem. Likewise, Fig. 9, which lacks 
a meristem as yet, may be on the pathway to more normal growth 
or is becoming ameristic. 

From the original group of gametophytes, a very few grew on to 
become mature, normal-appearing thalli. Ihese few were generally 
those whose development was most nearly normal. Thus among 
the survivors, a cordate thallus was sometimes encountered 
{Fig. 13). These tended to retain a rather shallow sinus. At the 
other extreme of variation were highly irregular, lobed forms 
{Fig. 14), The remainder were intermediate. Both types of game- 
tangia were produced and, at about one year, a few sporophytes 
appeared. These were apparently the products of fertilization by 
functional gametes. Several sporophji;es survived for some time. 
They were rather similar to one another and more closely re- 

American Fern Journal 

Volume 61, Plate 19 


Figs. 13-17. Pellaea uRiDGBsu X P. mucronata. Fios. 13, 15 from 
Pray 1858. (USC). Figs. 14, 16, 17 from Pray 1861 (USC). Fig. 14 is a 


Gametophytes of Natural Hybrids in Pellaea 133 

sembled the hybrid plants than they did either parental species. 
This lack of variability suggests apogamy. No evidence of asexual 
buds was noted however. All of. the young sporophytes were 
eventually lost while they were still immature. 



It might be expected that in view of tlie intermediate nature of 
the hybrid spbrophyte that the gametophytic generation would 
also show a compromise of the difTerences between the probable 
parents. The gametophytes of the parental species are markedly 
distinct in developmental features (Pray, 196Sa). While the 
mature thalhis of both species is cordate, the younger stages are 
distinguishable. In P. mucronata there is the regular difierentiation 
of a strongly lateral meristem which only gradually becomes 
shifted to a terminal and eventually median position on the 
maturing thallus. In P. hrkhesii, on the other hand, the marginal 
meristem is terminal from the beginning or at most only slightly 
asymmetrically situated; no truly lateral meristems occur. Thus 
it is the original site of the meristem which most obviously dis- 
tinguishes the gametophytes of the two species. In the hybrid, 
however, the great majority of the thalli never differentiate a 
meristem of any kind. Of those few which do, some are similar to 
one parent, some more like the other. This situation may indeed be 
a compromise between the parental pathways of development. It 
is of interest to note that certain Pellaea species, such as P. term- 
folia, are also characterized by just such variability in the site of 
meristem differentiation (Pray, 1968a). On the other hand, the 
filamentous or amcristic thalli are unlike anything shown by 
either parent. 

There is- also the question as to what extent the data from this 
particular hybrid are typical of nearly sterile fern hybrids. On this 
point, some observations on an other example are pertinent. A 
specimen was collected by the author in Arizona which appeared 
to be P. longiynucronala {Pray 3218),. ISIore careful examination 
suggests that the plant is actually the hybrid of P. lomjimucronala 
and P. wrujhliana. The specimen showed a high degree of sporangial 

American Fern Journal 

Volume 61, Plate 20 





I nun 


Figs. 18-29. Pellaea longimucronata X P. wrigutiana from Pray 

3218 (USC). 

Gametophites of Natural Hvbrids in Pellaea 135 

abortion but did have a few apparently good spores. When the 
spores were sown on agar, a few gametophytes did develop which 
exhibited the same variability and high frequency of aberrant 
types (Figs. 18-29) as described here for the California plants. A 
few thalli (Figs, 18 ^ 19, 23) had developmental sequences much 
Hke that of the parental species (Pray, 19G8a). Filamentous 
{Figs. 20, 25, 26) and highly variable elongated forms {Figs. 21, 22, 
24, 27-29) were also present. Ihe great majority of thalli were 
thus quite different from what might be expected. The most 
obvious effect of hybridity was the extension of the filamentous 
phase and the inability to initiate normal two dimensional growth 
and marginal meristems. It would appear then' that the preceeding 
account does not constitute an isolated instance, but rather that 
similar aberrant growth patterns are to be expected from the 
gametophytes of hybrids of this type. 

It may be significant that previous examples of gametophytes 
which closely resemble those of these nearly sterile hybrids have 
been recorded for certain apogamous ferns. Thus the thalli of 
some apogamous samples of P. andromedaefoUa (Pray, 1968b) 
closely match the more peculiar forms of the hybrids described 
above. Other apogamous Pellaeas (Pray, 1970) may also have 
similar forms. On the other hand, that apogamy is not necessarily 
associated with aberrant gametophytic development is also clear. 
Again, in the case of P. andromedaefGlia one apogamous popula- 
tion had quite regular gametophytes. Such exceptions were also 
noted in some other asexual Pellaeas, P. atroptirpiirea for example 
(Pray, 1968a). In view of the resemblances between the hybrid 
gametophytes and those of many apogams, it is noteworthy that 
apogamy is often associated with taxa or ''races" which have 
arisen through hybridization (Tryon, 1968). The difference is, of 
course, that these latter have become stabilized and self-per- 
petuating through apogamy, whereas the hybrids described m 
this paper apparently have not done so (as yet?). 

Finally, one might wonder whether the variations and irregulari- 
ties of development in the gametophyte generation of these hybrid 
ferns have the same basis as, or are analogous to, what Wagner 

136 American Fern Journal 

(1962) has discussed in the case of the sporophytic features of 
hybrid ferns. For the hybrids described by Wagner there was 
frequent asymmetry and irregularity of various leaf characteristics 
by which the parental species differed. Particularly pertinent here ' 
was the conclusion that ''hybrid expression is not a vuiiform and 
regular compromise." While this statement also applies to the 
situation described in this paper for ganif^tophytes, the causes 


may not be the same. In the sporophytes full genomes from the 
parental species are present and interacting. In the spores which 
produce the hybrid gametophytes such is probably not the case. 
Without a thorough investigation of sporogenesis and chromo- 
somal assortment in the hybrids, the chromosomal complements 
present in the various gametophytes remains unknown. It is, in 
fact, likely that they are variable chromosomally unless the 
functional spores are all the result of a non-reductional process. 
Chromosomal variability might explain the extreme variability of 
the gametophytes. 



stiped series. Phytomorphology 18: 113-143. 

. 1968b. Interpopulational variation in the gametophytes of 

Pellaea andromedaefolia. Amer. J. Bot. 55: 951-960. 
. 1970, The gametophytes of Pellaea section Pellaea: Light-stiped 

series. Phj^tomorphology 20: 137-144. 
Tryon, a. F. 1968. Comparison of sexual and apogamous races in the fern 

genus Pellaea. Rhodora 70: 1-24. 
Wagner, W. H., Jr. 1962. Irregular morphological development in hybrid 

ferns. Phytomorphology 12: 87-100. 

Department of Biological Sciexces, University of South- 
ern California, Los Angeles, CA 90007. 


Schemochromic Blue Leaf -surfaces of Selaginella 

Denis L. Fox^ and James R. Wells 

Schemochromic (i.e., optical or structural, non-pigmeiitary) 
colors are common in many animals, e.g., (1) Tyndall-blue scatter- 
ing in the iris of blue eyes; in skin areas of some birds, fishes, 
reptiles, and primates; in many feathers, as in jays; and (2) inter- 
ference-produced iridescence in pearl, nacre, wing-covers of some 
beetles, wing-scales of Morpho and many other butterflies, in 

feathers of peafowl, Hummingbirds, pigeons, and other birds 
(Fox, 1953). 

Although plants excel animals in the numbers and chemical 
varieties of true biochromes (colored molecules or pigments) that 
they store, instances of schemochromic coloration exhibited by 
plants are very rare. Moreover, the few examples involve no 
Tyndall-blue scattering, but are limited to the display of change- 
able spectral interference-colors, notably by a few seaweed species 
such as IridophycuSy Chondria, Zonaria, and others as viewed 
immersed in sea-water. A few exceptions found among land 
plants include some LycGpcdium species^ and certain Selaginella 
species, e.g., S. itilldenovii (Desv.) Paker, a native of ^Malaysia.^ 
The latter species has been introduced elsewhere, and is main- 
tained in greenhouses, including the University of Michigan's 
jMatthaei Botanical Garden, where the specimens studied were 
obtained through the kindness of Professor W. H, Wagner, Jr. 

^Distinguished Scholar, Cranbrook Institute of Science, 1970-1971; per- 
manent address: Division of Marine Biology, Scripps Institution of Oceanog- 
raphy, University of California, San Diego; La Jolla, CA 92037. I thank Dr. 
Warren L. Wittry, Director, and his colleagues at the Institute, for the hpnor 
and the many courtesies extended during a year's appointnient there. 

^ C. V. Morton and I. L. Wiggins have reported independently to us their 
observations of surface-iridescence on pinnae of some species of true ferns. 

^ There are also a few true ferns that exhibit iridescence on the adaxial 
surfaces, such as Anemia makrinii Maxon; hoMever, in this species at least 
this disappears after drying.— C.V.M. 

138 American Fern Journal 


The upper surfaces of this plant's leaflets exhibit, by sunlight 
or diffuse light, striking, changeable blue colors, reminiscent of 
the wings of Morpho butterflies but less brilliant. ]\Iicroscopic 
inspection by reflected light, under medium powers, reveals count- 
less microspheric raised areoles, like masses of very minute beads, 
highlighted on the upper, projecting surfaces and exhibiting 
successive blue, blue-green, and purple or violet hues, depending 
on the increasing angle of vision. When viewed at a fairly acute 
angle, the leaf's optical effects vanish, the surfaces then exhibit 
merely the green color of chlorophyll. 

It has been somewhat surprising not to encounter more pub- 
lished references to these striking changeable chromatic features 
over the upper leaflet surfaces of Selaginella. Walton and Alston 
(1938) commented upon the '^metallic sheen of their leaves • . , due 
to the presence of particles in. the cutin of the epidermis which 
reflect certain rays of light." 

While those authors came close to the factual basis of the 
observed coloration, the reference to '^particles" was perhaps 
misleading, connoting as it does the presence of substantial, per- 
sistent material, perhaps of a solid character. The microsurfaces 
which reflect a changeable spectrum of optical colors are in fact 
air which is distributed into a pattern of minute films; hence the 
air is not discharged, nor the color quenched, by squeezing or 
rubbing a leaflet between the fingers. 

The optical colors disappear when the leaves are allowed to 
wilt. They are discharged also on wetting the fresh leaves briefly 
with either water or ethanol, but on evaporation or blotting of the 
fluid, the colors are restored. However, if a wetting agent, e.g., 
the clearing fluid Aerosol (Eimer and Amend) is applied, the 
immediate quenching of the bright colors is permanent, even after 
the treated specimen is rinsed in water and superficially dried. 

Examination by transmitted light under high optical powers 
(400 X) reveals stomates oa er the aba^ial leaf-surfaces and many 
small, light-refracting, air-filled spaces on the adaxial surfaces. 
The individual dark, light-refracting elements become suddenly 
clear as water gradually fills the air-spaces. 

ScHEMocHROMic Leaf-suhfaces OF Selagixella 


The exposed cuticular layer of air-laden micro-spaces between 
the regular, projecting elements of the bead-like clusters, thus 
reminiscent of very minute interstitial soap-films^ is the seat of the 


iridescent color-production. In much the same manner, entrapped 
microfilms of air give rise to the striking changeable blue colors of 
Morpho wings, which likewise are instantly quenched by the 
addition of foreign fluids having the same refractive index as the 
solid wing-protein, and which are restored on leaching away the 
fluid or allowing it to evaporate (Fox, 1953): 

Whether this unusual chromogenic manner of storing air on the 
upper surfaces of the leaflets may be merely an incidental condi- 
tion, evoked by photosynthetic, respirational and transpirational 
processes, and may be of no selective advantage to the plant, or 
whether it may in fact serve a role of physiological or biophysical 
importance (e.g., such as thermal buffering, light-screening, or 
water-retaining functions) must remain a problem for plant 

Literature Cited 
Fox, D. L. 1953. Animal Biochromes and Structural Colours. Cambridge 

University Press. 



/"II: Lycopodiinae. 
Martinus Nijhoff, 


Cranbrook Institute of Science, Bloomfield Hills, ^II 

140 American Fern Journal 

Shorter Notes 


Asiatic Fern.— In the "Index Filicum'' of .1905, Asplenium 
kamfchatkannm Gilbert (Fern Bull. 9 : 54. 1901) is listed as dubious, 
with the comment ''Athyrium sp.?" Apparently no one has ever 
examined the type, which is right here in the U, S. National 
Museum. It was collected on the U. S, North Pacific Exploring 
Expedition on the Ochotsk Sea, Kamtchatka, by J. Small, be- 
tween 1853 and 1850, aiid had been originally identified as Asvlen- 
ium fdix-femina L. The same species was collected also by the 
same expedition on Yezo Island. 

This species was overlooked in Komarov's ''Flora of Kamt- 
chatka'' (1927), and in Hulten's "Flora of Kamtchatka and Ad- 
jacent Islands" (1930) it is mentioned only as an unknown species 
of doubtful position. The type shows that this plant is the same as 
Athyrium pteroraciiis Christ, a species reported in Ohwi's "Flora 
of Japan'' from Hokkaido, Honshu, Sakhahn, southern Kurile 
Islands, and Kamtchatka. It is a rather distinctive species, 
especially in the pubescent costae beneath, and clearly has nothing 
to do with Athyrhim^ fdix-Jemina. In fact, this species and some 
others have in recent years been removed entirely from Athynum 
and placed in Dryoathyriujn, Lunathyrium, or Par athyrium. If 
one assumes that the rachis structure is of fundamental importance 
in this group of ferns, then indeed this species and its allies will 
have to be separated from Athyrium^ which differs in having the 
rhachis groove above interrupted and open to receive the grooves 
of the lateral pinnae. The matter has been discussed most re- 
cently by Hideaki Ohba in his paper "Considerations on the Genus 
Lunathyrium of Japan'' (Sci. Rep. Yokosuka City Mus. 11 ^ 
48-55. 1965). He recognizes the single genus Lunathyrium Koidz. 
(1932) Avith three sections — sect. Lunathyrium, sect. Athyriopsis 
{Athyriopsis Ching, Acta Phytotax. Sinica 9: 63. 1964; Type: 
L. japonicum)^ and sect. Dryoathyrium (Dryoathyrium Ching, 
Bull Fan Mem. Inst. Biol. Bot. 11:79. 1941; Type: L. horyanum; 
a synonym is Parathyrium Holttum, 1958). This seems the most 

Shorter Notes " 141 

reasonable treatment. The essential synonymy is as follows: 

LuNATHYRiUM PTEKORACHis (Christ) Kurutu in Xame^ala & Kuratu, Enum. 

Jap. Pterid. .309. 1961. [reprint without change of paging from 

'* Collection and Cultivation of Our Ferns and Fern Allies" by T. 

Aikyrium pterorachis Christ, Bull. Herb. Boiss. 4: 668. 1896. 
Asphnium kamtchatkanuyn Gilbert, Fern Bull. 9; 54. 1901. 
Asplenium kamlchulicmn Hulten, Kungl. Sv. Vet. Akad. Handl. 8(2): 237. 

1930. Unintentional and illegitimate renaming of A. kamtrhatkanum 



. DryoaUiyrium plerorachid Ching, Bull. Fan Mem. Inst. Biol. Bot. 11: 81. 

1941 (as "pterachis" erroneouslj-). 
Pamthyrium pterorachis Holttum, Kew Bull. 1958: 449. 1959. 

C. V. INIoRTON, National Museum of Xatnral History, Washing- 
ton, DC 20560. 

A New Locality for Pbilotu.m nudum in Sonora, IMexicg — 
In the spring of 1970, while studying the palm oases on the central 
coast of western Sonora, ^Mexico, I discovered a small colony of 
Psilotum nudum Palisot in Xacapule Canyon in the mountains 
near San Carlos Pay and about 10 miles northwest of Guaymas. 
The mouth of this canyon is located about 4.5 miles in from the 
coast, and the narrow canyon drains to the east out of steep and 
rugged volcanic mountains. The Psilotum grew at an alga-covered 
seep on the canyon wall in the shade of palms, W ashingtoni a ro- 
husta Wendl., and a few trees of a fig, Ficus padifdia H.B.K. While 
this locality is definitely in the Sonoran Desert, the shade of the 
canyon and the constant water supply provided by springs and 
seeps allows a number of plants with high water requirements to 
survive. Only one colony of Psilotum with about a dozen stems 
two to three inches high was found. A small collection of this 
plant (Boutin & Brandt 2814) is now growing in the Huntington 

Botanical Gardens. 

Besides the Nacapule Canyon locality, three other stations are 
known for P. nudum in southern Arizona, Sonora, and western 

142 American Fern Journal 

Chihuahua. It was recently reported in Sycamore Canyon, Arizona 
by Phillips (Amer. Fern J. 60: 30-32. 1970). It is also known in 
the Sierra Saguaribo of eastern Sonora in the drainage of the Rio 
Mayo, as reported by Gentry (Rio Mayo Plants, 1942), and from 
the Barranca del Cobre of Chihuahua, reported by Knobloch and 
Correll (Ferns and Fern Allies of Chihuahua, 1962) and also re- 


cently collected by Kimnach and Brandt (980).' The habitat of 
Knobloch's collection w^as not reported; however, each of the 
other collections has been recorded as being from a rock crevice 


or cliff site. — Fredrick C. Boutin, Huntin^tcji Botanical Gar- 
dens, San Marino, CA 91108. 

Recent Fern Literature 

The Classification of the Cyatheaceae, by R. M. Tryon, 
Jr. Contr. Gray Herb. 200: 4-53. 1970.— The scaly Cyatheaceae 
have mostly been divided into three traditional genera — Cyathea, 
with a complete indusium, Hemitelia, with a lateral, semispherical 
indusium, and Alsophila, with the indusium reduced to a small 
scale or absent. It has seemed to many that these three genera, 
although admittedly rather artificial, do represent natural groups, 
at least so far as the American species are concerned, for it has 
seemed that the species were more closely related to others in their 
own group than to those of other groups. However, Copeland, 
followed by Domin, did not agree and lumped all the. species in a 
single large genus, without any subgeneric classification or other 
grouping. Recently, Holttum, working on Old World species, 


established a new classification in which the scale characters were 
regarded as more important than the indusial characters. Tryon 
has followed Holttum by working on the New World species, and 
has developed a new system. He divides Cyathea into six genera — 
Sphaeropteris, Alsophila, Nephelea, Trichipteris, Cyathea, and 
C neniidar ia . Sphaeropteris, mostly Old World, contains those 
species with the cells of the petiolar scales all alike. The other 
genera have the marginal cells variously differentiated. Alsophila 
and Nephelea have the petiolar scales with a dark apical seta, and 

Recent Fern Literatuue 143 


sometimes also such setae on the back or margins. AhopJiila^ also 
mostly Old World, has no spines or, if these are present, they are 
not "corticinate". Nephelea, strictly New World, has '^squaminate" 
spines, the smaller ones being thin-marghiate and scalclike. 
Cnemidariaj Cyathea, and Trichipteris have the petiolar scales 
lacking setae. In Cnemidariaj strictly American, tlie veins are 
areolate or, if free, the basal veins on each side are connivent to 
the sinus. Cyathea and Trichiptens have the veins all free and 
reaching the margin above the sinus. Trichipteris^^ strictly Ameri- 
can, includes most of the exindusiate species that have been called 
Alsophila, Cyathea, also strictly American as here delimited, 
includes the indusiate species. In addition to the genera mentioned 
above, the generally recognized genera Lophosoria and Meiaxya 
are treated. 


Tryon believes his classification to be natural. It may be so, 
although it does have the appearance of being a ' 'one-character" 
classification, the scale character being the decisive one in placing 
a species into a genus. It will certainly be useful in identifying 
specimens, and the stressing of the petiolar scale characters may 
influence collectors to be more careful to collect full specimens 
rather than detached pinnae. Tryon has not really justified his 
recognization of these groups as mdependent genera, his only 
statement being that he believes each genus to be a major evolu- 
tionary line, but that could and should be equally true also of 
subgenera. However that may be, Tryon is to be congratulated 
on a major contribution to the taxonomy of the Cyatheaceae. He 
is likely to be widely followed. — C.V.IM. 

are an 

^ It is unfortunate that Tryon adopts the spelling ''Trichipteris:' He states 
that the original spelHng must stand. This is not true by the Code. Recom- 
mendation 73G, d, indicates that the proper connecting vowel in Greek in 
cases like this is an "o.'' Although this is only a recommendation, it is also 
made into a Rvile by Art. 73, Note 2, which states that a wrong connecting 
vowel is treated as an orthographic error; the "Notes'' in each article a: 
integral part of the Rules, put in to explain the operation of the Rule itself. 
The main body of Art. 73 indicates that orthographic errors are to be cor- 
rected. Therefore, the correct name must be Trichopteris. Since this is merely 
a correction rather than a different name it will continue to be credited to 
Presl in 1822. The matter is exactly like the use of a wrong connecting vowel in 
specific epithets, which also must be corrected by Art. 73. 

Hi American Fern Journal 


A Manual of Plant Names (Second revised edition), by C. 

Chicheley Plowden. pp. 1-260. 1970. Published by Philosophical 
Library, 15 East 40 St., New York, N. Y., 10016. $10.00.— This 
book is intended for the gardener and amateur botanist who is 
interested in the meaning of plant names. Those treated are pri- 
marily plants cultivated for food, drugs, or ornament, and the 
commoner wild plants. The etymology of the Latin generic and ' 
specific names is given, and also a glossary of botanical terms 
used in descriptive botany. Needless to say, it will not replace a 
scholarly and indispensible work like II. W. Brown's ^'Composition 
of Scientific Words" or B. D. Jackson's ''A Glossary of Botanic 
Terms," but it may find a place by those interested in less technical 
and less complete works. I would not myself completely trust the 
etymologies given (although I must confess that I have not checked 
many of them). For instance, Glechovia is said to be derived from 
'^glaukos" (gray), but it is surely from ''Glechon," the Greek 
name of some kind of pennyroyal. Pteris is said to be from ^'pteron, 
a wing/' but it is actually merely the Greek word pteris, applied to 
a fern; pteris may have come originally from ^^pteron," which 
means primarily ^'feather" rather than 'Sving." Frayana is not 
derived from ''fragrans" but is a Linnaean variant of '^fraga," 
the Roman name for the strawberry; it may have ultimately 
been derived from **fragrans" by the Romans. But no matter, 
the derivations are suggestive and doubtless correct in most 
instances. — C- V. ]\L 

Exotic awl Hardy Ferns 



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R!]iOM, Wlseonsia 54971 

Vol. 61 October-December, 1971 No. 4 

American Jfern fournal 


Published fay the 









The Genus Grammatosorus C. V. Mobton 146 

The Genus Selaginella in Baja California, Mexico Ika L. Wiggins 149 

Notes on the Ferns of Dominica and St. Vincent, II 

C. V, Mobton and David B. Lellinger 161 

Two New Tree Ferns from Costa Rica Lms Diego G<5mez 1 

Comparative Habitat Requirements for Spore Germination and Pro- 



Growth of Three Ferns in Southeastern Michigan 

Hill 171 

The Gametophyte of Thelypteris erubescens Lenette R. Atkinson 183 

Shorter Notes: A Recent Find of Isoetes in Louisiana; Jamesonia 

alstonii in Oaiaca, Mexico 186 

Recent Fern Literature 188 

Index to Volume 61 . i 188 


Mfssovm Bor*^"Tf«rc 


^ ' 

^'^M - I . ,0, JAN 10 1972 

(Cfie American Jfern ^ocietp 

Council for X971 

Warhen IL Wagner, Jr., Department of Botany, University of Michigan, 
Ann Arbor, Michigan 48104. President 

John T. Mtckel, New York Botanical Garden, Bronx Park, Bronx, New 
York. 10458. Vice-President 

Richard L. Hauke, Department of Botany, University of Rhode Island, 

Kingston, Rhode Island 02881 


LeRoy K. Henry, Division of Plants, Carnegie Museum, Pittsburgh, Penn- 
sylvania 15213. Treasurer 

David B. Lellinger, Smithsonian Institution, Washington, D. C. 20560. 


Rational ^ocietp 9^epres(entatibes 

Warren II. Wagner, Jr., University of Michigan AA.A.S. Representative 
Roll A M. Tryon, Jr., Harvard University AJ.B,S, Representative 

SImerican jfern SToucnal 


David B. Lellinger Smithsonian Institution, Washington, D. C. 20560. 

C. V. Morton Smithsonian Institution, Washington, D. C. 20560. 

Roll A M. Tryon, Jr. 

Gray Herbarium, Harvard University, Cambridge, Mass. 02138 

Ira L. Wiggins Dudley Herbarium, Stanford University, Stanford, Calif. 


An illustrated quarterly devoted to the general study of ferns, owned by 
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Changes of address, applications for membership, subscriptions, orders 
for back numbers should be addressed to the Treasurer. 

Subscriptions $5.50 gross, $5.00 net (agency fee $0.50); sent free to mem- 
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American Jfern Journal 

Vol. 61 October-December, 1971 No. 4 

The Genus Grammatosorus 

C. V. Morton 

The almost unknown species Gravwmtosoms blumemms Kegel 
was published in the "Index Seminum" of the botanical garden in 
St. Petersburg in 1866; it was based on a cultivated plant that 
Kegel had received from a man named Booth under the unpub- 
lished name of "Aspidium blumei." This Booth was probably 
John Booth, of Flottbeck, who according to the "Index Herbar- 
iorum Index of Collectors" collected in Java. Kegel indicated that 
the original habitat was probably the East Indies. The connection 

with Blume is obscure. 

J. INIilde obtained a specimen from Kegel and studied it care- 
fully, and published his results in the "Botanische Zeitung" (26: 
614-617. 1S68). He concluded that the plant might represent 
either a sport developing in cultivation or a hybrid. 

This plant has remained essentially unknown for the last hundred 
years. It disappeared rapidly from cultivation, and no wild speci- 
mens resembling it have been discovered. Through the courtesy of 
the Director, Botanical Institute of the Academy of Sciences of 
the U.S.S.K. in Leningrad, I have been permitted to have the 
type on loan; a photograph of it is given herewith. It is a rather 
poor specimen, lacking a rhizome, The habit and venation are 
those of rectaria. The general irregularity in cutting makes it 
appear that this may indeed be a hybrid, and this is partly con- 
firmed by the fact that most of the sporangia are abortive. A 
photograph of one of the larger and better developed spores made 
by the scanning electron microscope is shown in the accompanying 
figure, but even this is not a fully developed and regular spore. 

Volume 61, Number 3, of the Joubn.^l, pp. 97-144, was issued Nov. 15, 1971. 

American Fezin Journal 

Volume 61, Plate 21 





f ^ ^^^ f...l PHW^tt&f** ' ^ 

-^^ ^.. ^ • ^ .' 














n>' ^-^-'^ F ► ■ n FJ_JlL_>' J 


The Genijs Grammatosorus 


The chief pecuharity of Grammatosorus that caused Kegel to 
describe this as a new genus is the soruSj which is strongly elongate 
along the reticulate vcinlets; it is obviously indusiate throughout. 
Elongate^ indusiate sori such as might occur in Diplazhim are 

Scanning electron microscope photograph of Grammatosorus blu- 

MEANUS, X 570. 

not characteristic of Tedaria. There is however one species in 
which such sori occur, namely the extremely peculiar Tedaria 
dolichosora Copeland (PhiL J. Sci. 38: 136. 1929); in this the 
sori are strongly elongate and indusiate; they curve around at the 
tip, thus becoming athyrioid in shape, and come back down the 
outer side of the veinlet, sometimes clear to the bottom, thus 

148 American Fern Journal 

making a truly diplazioid, double sorus, with the indusia back to 
back and the sporangia spreading in both directions. In Grammai- 
osorus the sori do sometimes curve a little at the apex where there 
is a vein fork, but they do not come back on the other side to form 
a diplazioid sorus. Other than in the sori there is no obvious alliance 
between Grammato sorus and T. dolichosora. In the latter the blade 
is much more divided and the axes are dark as in T. melanocaulon 
(Blume) Copel.i In Grammatosorus, the stipe and rhachis are 

It may indeed be that Grammatosorus is a hybrid, in which case 
one of the parents might be Tedaria (Arcypteris) irregularis (K. 
B. Presl) Copel.; I would not dare guess as to the other parent, 
which would probably be in another genus. On the other hand, 
the existence of another species, T. dolichosora, with somewhat 
similar soriation indicates that this plant might well be a true 
Tectaria, for there is no evident reason to think that T. dolichosora 
is also a hybrid. In order to provide this plant with a nomen- 
claturally correct name, the following new combination is proposed: 

Tectaria blumeana (Kegel) Morton, comb, no v. 

Grammatosorus Uumeanus Kegel, Ind. Sem. Hort. Petrop. 1866: 75. 1866. 

Aspidium grammatosorus Salom. Nona. 54. 1883. A superfluous and illegiti- 
mate name, since Salomon ought to have adopted the epithet blumeanum, 
which was available. Doubtless he did not because of the existence of an 
Aspidium blumei Kunze ex Mett. (1856), an entirely different plant. 
However, according to the Code both blumei and blumeanum are possible 
and correct, since thay are considered different names. 

National Museum of Natural History, Washington, 
DC 20560. 

moll^'f f^"?^-'^ wrongly spelled "melanoraulis" by Copeland, apparently to 

^f Rf.l ■"'''""t\° ^F''^ ^^'^^ ^^'^taria. However, the original melanoraxdon 
ot Blume is a substantive and therefore invariable. 

Selaginella in Baja California 149 

The Genus Selaginella in Baja California, Mexico 

Ira L. Wiggins 

The genus Selaginella was omitted from the '^Vegetation and 
Flora of the Sonoran Desert" (Shreve and Wiggins, 1964) because 
all material available to me at the time had been collected at 
altitudes above the elevation we had accepted as marking the 
upper limit of the Sonoran Desert, or came from areas in the Upper 
Sonoran Zone in northwestern Baja California- But when one 
considers the entire peninsula of Baja California, it contains seven 
species of Spike-moss, five of which Tryon places in the subgenus 
Selaginella section Tetragonostachys (A. Braun) Hieron. (Ajin. j\Io. 
Bot. Gard. 42 : 1-99. 1955). The other two belong in the subgenus 
Stachygynandrum series Circinatae, according to Alston (in Ver- 
doorn, '^Manual of Pteridology," pp. 503-4. 1938), These seven 
Spike-mosses may be distinguished with the aid of the following 
key, and are illustrated here by line drawings in Plate 23 ei seq. 
The drawings are offered as supplements to the excellent half-tone 
reproductions in Tryon's paper in order to emphasize certain 

Key to the Species of Selaginella in Baja Californlv 

1. Stems (with attached leaves) terete or quadrangular, not dorsiventrally 

flattened; plants erect, ascending, or decumbent but not forming rosettes, 

not rolling into a ball when dr}'; leaves narrowly lanceolate to linear-Ianceo- 

late, with a median longitudinal groove on abaxial side (Subg. Selaginella). 

2. Stems erect or ascending, not decumbent or prostrate; rhizophores 

produced only at or near bases of main stems (Ser. Arenicolae) . 

3. *S, bigelovii 

2. Stems creeping or prostrate, the main ones neither erect nor strongly 
ascending; rhizophores along lower side of most of stem. 

3. Setae at apex of vegetative leaves and sporophylls denticulate 
scabrous ; leaves markedly glaucous ; sporophylls acuminate 

(Ser. Rupcstres) 2. *S. aspreUa 

3, Setae at apex of leaves smooth, filiform and ephemeral, or lacking; 
leaves bright green or persistently ashy-gray (in *S. cinerascens) , 
not glaucous. 


AN FeRiN Jol'rxal 

Volume 61, Plate 22 

Distribution of Selagixella in Baja California. Solid circle = S. 
arizonica; Open circle = S. aspre-lla; Open triangle = S. bigelovix; 
Solid triangle - S. cinerascens; Open square = S. eremophila; Solid 
square = S. lepidophylla; Solid star = S. pallescens. 

Selaginella in Baja California 151 

4. Ultimate branohlets mostly 2-4 mm long, these and main 
stems appressed-prostrate; leaves ashy-gray, their tips acute 
or sometimes muticous, not setiferousj plants growing on 

soil, not in rock crevices (Ser. Sartorii)., 4. S. cm€ra,'iccns 

4. Ultimate branches mostly 5-12 mm long; tip« of branches 
turned upward, not tightly appressed-prostrate; leaves bright 
green, not ashy, their tips bearing straight, slender, more or 
less deciduous setae, or the terminal seta (in *S'. eremophila) 
filiform and ephemeral, usually visible only on leaves at tips 
of branchlets (Ser. Eremophilae). 

5. Microspores usually orange; terminal seta straight, 
milk-white, 0.1-0.3 mm long, never equaling leaf blade, 

often deciduous in age \. S, arizonica 

5. Microspores usually yellow; terminal seta filamentous, 
tortuous, often one-half to equaling length of blade, 
deciduous very early and obvious only on leaves at tips 
of growing branchlets 5. S. eremophila 

1. Stems and branches flattened dorsiventrally; plants prostrate and rosu- 
late when moist, branches curling inward and upward to form a compact 
ball when dry; leaves dimorphic, those on upper side of stem smaller than 
ones on low^er side, neither series longitudinally grooved (Subg. Stachy- 
gynandriun Ser. Circinaiae), 

6. Leaves on lower side of stem suborbicular, rounded at apex, reddish 
or brownish; leaves on upper side of stem broadly ovate, without a 

terminal seta, conspicuously white-margined 6. ^S. lepidophylla 

6. Leaves on low^er side of stem ovate, rarely suborbicular, silvery white, 
acute and short-setiferous at apex; leaves on upper side of stem ovate 
to orbicular, bearing a stout, terminal, denticulate-scabrous seta, less 
markedly white-margined 7. 5. paUesccns 

Descriptions published by Maxon (Smiths. i\Iisc. Coll. 72(5) : 
1-10, ph. 1-6. 1920) and by Weatherby (Amer. Fern J. 33: 113- 
119. 1943; J. Arn. Arb. 25: 407-419, p/^. 1, 2. 1944) admirably 
cover the species within the subgenus Selagiriella that are known to 
occur in Paja California. Likewise^ those in Correirs 'Terns and 
Fern Allies of Texas," pp. 28-43. 19o6, and in Knobloch and 
Correll's 'Terns and Fern Alhes of Chihuahua/' pp. 16-32. 19G2, 
deal with some of the species in subgenus Selaginella and both of 
the species in subgenus Stachygynandnim. Therefore, I include 
here only citations of specimens from Paja California examined, 
together with comments on habitats and distribution. 

American Fern 

Volume 61, Plate 23 

Selaginella in Baja California 153 

The line drawings were made with the aid of a Spencer camera 
lucida and various lens combinations of a Eausch and Lomb 
compound microscope. 

Abbreviations of herbaria from which specimens were studied 
are those recommended by Lanjouw and Stafleu in ^^Index Her- 
bariorum," ed. 5, 1964. 

1. Selaginella arizonica Maxon, Smiths. Misc. Coll. 72(5); 5, 

pi 3. 1920. 
On rock faces, in crevices in and between boulders and ledges, 
often on soil or gravel at the bases of boulders, but rarely away from 
boulders or cliff faces, in Gila, Graham, Maricopa, Pima, and Pinal 
counties, Arizona, east in southwestern Texas (although not known 
to occur in New Mexico), south into northern Sonora, and known 
from a single collection near Loreto, Baja California. Mostly at 
600 to 2000 m alt. 

Specimens examined: Baja California: Arroyo Undo [Hondo], [near] 
Loreto, 26 Oct 1930, M. E. Jones (DS). 

The disjunct distribution of this Selaginella is puzzling, for there 
are areas along the eastern slopes of the Sierra San Pedro Martir 
and other ranges in Baja California that would appear to provide 
suitable habitats for it. More intensive search may reveal it in 
such areas. 

2. Selaginella asprella Maxon, Smiths. ]VIisc. Coll. 72(5) ; 6, 

j)l 4' 1920. 
On rock faces, in crevices on ledges, at bases of boulders, cliffs, 
and around talus, Los Angeles, Orange, and San Bernardino 

Figs, a-e, Selaginella ahjzonica: a = Functional leaf from under 


SPOROPHYLL, abaxial SIDE. FiGS. f^i, Sel.\ginella asprella: f = Leaf from 


opposite f; h = Megasporophyll, abaxial sLTtFACE; i = Mature micro- 
spore. Figs, k-p, Selaginella bigelovii: k = Mature megaspore, outer 
surface; 1 = Microspore, commissural side; m = Microspore, outer 
surface; n, o = Leaves from same level on opposite sides of ascending 
stem; p = Abaxial view of megasporophyll. 

154 Ajmerican Fern Journal 

countieSj California, south along the eastern escarpment of the 
San Jacinto and Cuyamaca Mountains into Baja California. 

Specimens examined: Baja California: In crevices of granitic rocks along 
streams, near La Encantada, Sierra San Pedro Mdrtir, alt 7800 ft, 22 Sept 
1930, Wiggins <fc Demaree 5002 (DS) ; Rocky cliff in first canyon off Canon del 
Diablo on NW side of Picacho del Diablo, eastern flank of Sierra San Pedro 
M^rtir, alt 6000 ft, 15 June 1954, Chambers 64S (DS); Rim overlooking canon 
between main range and Picacho del Diablo (Picacho de la Providencia), near 
Lat. 31° N, Long. 115"^ 24' W, about 9000 ft, 5 Sept 1961, Wiggins 16635 (DS) ; 
Crevices in granitic rocks on SE slopes, Cerro Venado Blanco, Sierra San 
Pedro Martir, 15 Sept 196S, Moran 15670 (CAS). 

This species is far more common among igneous rocks and 
cliffs in the Sierra San Pedro Martir than the small number of 
specimens in herbaria would indicate. 

3. Selaginella BiGELovii Underw. Bull. Torrey Bot. Club 25: 

130. 1898. 

On dry, sunny, rocky slopes and ridges or on soil in partial shade 
of sparse stands of chaparral, often abundant around boulders or 
on steeply sloping to nearly vertical cliffs and rock faces, inner 
Coast Ranges, Santa Clara County, California, southward to the 
mountains of the Cape Region, Baja California, at elevations 
between sea level and ir)00 feet in the northern part of its range, 
and upward to over 6000 feet in the Cape Region. 

Specimens examined: Baja California del Norte ; Brush-covered hills SW of 
Tijuana on Agua Caliente road, 24 Feb 1934, R. S. Fern's 8470 (DS) ; South 
facing slope about 11.4 mi E of Rodriguez Dam, east of Tijuana, 24 Jan 1966, 
Wiggins 20373 (DS); Near Rodriguez Dam, 10 mi E of Tijuana, in tufts at 
bases of rocks, 1 Apr 1947, //. F. Copeland (DS) ; Hillsides NE of Ensenada, 
alt 200-400 ft, 23 Feb 1930, Wiggins 4212 (CAS, DS); About rocks on hill- 
sides facing the sea, Ensenada, 7 Apr 1921, /. M. Johnston 3005 (CAS); San 
Carlos River (S of Ensenada), 10 Sept 1923, Eastwood 12391 (CAS); San Car- 
los canyon, 10.6 mi E of Ensenada-Santo Tomas road, 8 Apr 1954, Wiggins 
13058 (DSj ; On dry soil and in crevices in rocks, hillsides above Arroyo de la 
Agua'Marga, southern Sierra San Pedro Mdrtir, alt ca 2700 ft, 13 May 1941, 
Wiggins 9937 (DS; ; Slopes of San Antonio Mesa, 11 April 1936, Epimg & 
Stewart (DSj ; About 8 mi E of El Aguajito along road from El Rosario to 
San Augustin, Lat. 30" 04' N, Long. 115° 21' W, alt ca 1500 ft, 6 Feb 1962, 
Wiggins tf- Thomas 68 (DS) ; About 7 mi S of main San Augustin-Laguna 
Chapala road, along track to Cerro Blanco and San Jose, Lat. 29° 42' N, 
Long. 114° 25' W, alt ca 1500 ft, 7 Feb 1962, Wiggins & Thomas 120 (DS); 

Selaginella in Baja California 155 

Catavind, 14 Apr 1954, Dorothy R. Harveij (CAS); North-facing rocky wall of 
canyon, 2 mi N of Mision de San Borja, 16 May 1959, /. L. & D. B. Wiggins 
14843 (CAS, DS); Rocky ridge above Rancho Aguajito, 32.5 mi SE of El 
Rosario, 15 May 1956, Howell 31012 (CAS); Same area. Porter 162 (CAS); 
Rocky talus slopes 5 mi W of Mision de San Borja, along road to Rosarito, 
alt ca 1000 ft, 18 Feb 1962, /. L. & D. B. Wiggins 16739 (DS); Between 
boulders in arroyo bottom opposite store at El Arenoso, alt ca 2500 ft, 21 Jan 
1960, Porter 162 (DS); On exposed rocks E slope of Cerro Barranco, Sierra 
Santa Lucia, near Lat. 26° 56' N, Long. 112° 28' W, alt ca 1300 m, 10 Mar 1964, 
Moran 11784 (DS); Locally common on north slopes near summit, Cerro San 
Juan, near Lat. 27° 58' N, Long. 113° W, alt ca 1200 m, 4 Feb 1964, Moran 
11697 (DS); NW slope, Cerro de la Mina de San Juan, near Lat. 28° 43' N, 
Long. 113° 38' W, 25 Mar 1960, Moran 8051 (DS). Territorio Baja California 
del Sur: Rocky mountain top, in oak forest, Cerro de la Giganta, alt 5000 ft 
plus, 1 Mar 1939, Gentry 4274 (DS) ; On steep, north-facing slope on ridge NW 
of main peak of Cerro de la Giganta, alt 1200-1350 m, Lat. 26° 8' N, Long. 
111° 36' W, 25 Nov 1947, Carter, Alexander <?' KvUogg 2050 (DS); Among 
granite rocks at summit, in Quercus idonea with occasional Q. devia and Pinus 
cenibroides, Sierra de la Laguna E of Todos Santos, alt ca 2000 m, 26 Dec 1947, 
Carter, Alexander d' Kellogg 2378 (DSj; In granitic rocks at edge of Arroyo del 
Leon, N of road from La Paz to Los Planes, 22 mi SE of La Paz, alt ca 1565 ft, 
11 Dec 1959, Wiggins, Carter & Ernst 494 (DS). 

This is one of the most widely ranging species among the species 
of Selayinella native to western North America, growing well 
through nearly 15 degrees of latitude, but mostly on the coastal 
slopes of the foothills and mountains within its range. It is also 
remarkable for the lack of variation between and within popula- 
tions, and for the diversity of the substrata it occupies. It seems to 
be equally adapted to granitic rocks, to basaltic extrusions, on 
sandstone, in crevices in serpentine, or gravel]}- detritus derived 
from any or all of these basic sources, and even on clay soil where 
little rock is present. It occurs too, on both north- and south- 
facing rocky areas if it is within the influence of coastal fog drift, 
and does well on vertical clifTs, gentle slopes or nearly level areas 
in shallow depressions. 

4. Selaginella cinerascens A. A. Eaton, Fern Bulk 7: 33. 1899. 
Forming close coverings of clay soil in open areas or among 
shrubs in coastal areas near San Diego and southward into Baja 

American Fern Journal 

Volume 61, Plate 24 

Selaginella in Baja California 157 

Specimen!S examined: Baja California: Among chaparral about 20 mi N of 
Ensenada, along road to Tijuana, 28 Feb 1934, Ferris 8474 (DS); Ensenada, 
mostly on hillsides, 7 Apr 1921, /. M. Johnston 3004 (CAS) ; about 30 mi S of 
Tijuana, 13 Mar 1956, Howell 30937 (CAS) ; HiUside NE of town of Ensenada, 
alt 200-400 ft, 23 Feb 1930, Wiggins 4213 (DS); about 12 mi N of Santo 
Tomds, along highway, 29 June 1962, Wiggins & Thomas 4I6 (DS). 

The paucity of specimens of this species in herbaria is doubtless 
attributable to the difficulty one has in freeing good specimens 
from the heavy soil on which it grows. When the substrate is moist 
it requires tedious, time-consuming effort to get the plants separ- 
ated from the clay, and when dry the earth is so hard that the 
fragile, closely attached plants are badly broken in the process. 

Consequently, most collectors ignore the close mats of moss-like 

The wide differences in the sizes of leaves found on a plant, as 

pointed out by Tryon, is not applicable to the leaves taken on the 

upper and lower surfaces of the stem or branches at the same 

distance from the apex (or base) of a stem or branch, as indicated 

by Plate 24, figs, a-c, 

5. Selaginella ereimophila Maxon, Smiths. Misc. ColL 72(5) : 3, 

pi 2. 1920. 
On shaded sides of rocks in open desert, along ledges and on 
steep to vertical walls of canyons, and occasionally on gravel at 

Figs, a-g, Selaginella cinerascens: a = Leaf from under side of 

SPORE, OUTER FACE. FiGS. h-1, Selaginella eremopiiila: h = Megaspore, 
OUTER face; i ^ Leaf from upper side of decumbent stem, terminal seta 
betached; k = Leaf from under side of stem, opposite i, terminal seta 
detached; 1 = Megasporophyll, ABAXIAL VIEW. FiGS. m-t, Selaginella 
lepidophylla: m = Tip of br.\nchlet viewed from upper smE; n = Seg- 



American Fern Journal 

the bases of such rocky areas, mostly below 3,000 feet, western 
Colorado Desert in California, in the Chuckwalla Mountains, 

Figs, a-c, Selaginella eremophila: a - Nearly fully developed leaf 



let; e = Leaf from upper side of branchlet; f 





1 1 



south in Baja California about to the vicinity of Eahia de los 



Mexicali, along highway to Alaska and Tijuana, alt ca 2500 ft, 6 Apr 1954, 

Selaginella in Baja California 159 

Wiggins 13045 (DS); Rocky hillside 0.5 mi SW of Rosarito, ca 40 mi SE of 
San Quintin, SW foothills of Sierra San Pedro Martir, alt ca 2900 ft, 16 May 
1944, Wiggins 9981 (DS) ; Lower half of Canon Diablo, east face of Sierra San 
Pedro Mdrtir, near Lat. 31° 2' N, Long. 115° 25' W, 22-27 Mar 1954, Stombler 
131 (DS); El Terminal, 16 mi S of Los Angeles Bay, common on N side of 
rocks near canyon mouth, near Lat. 28° 45' N, Long. 113° 35' W, alt ca 400 m, 
14 Jan 1962, Moran 8524 (DS) ; About 6 mi S of Las Flores, S of Los Angeles 
Bay, near Lat. 28° 43' N, Long 113° 32' W, 13 Feb 1962, Wiggins & Thomas 

The terminal seta on vegetative leaves of this species is re- 
markably slender and tortupus. However, this seta is very easily 
detached, so one rarely finds a leaf bearing a terminal seta except 
among the incurled, partially expanded young leaves at the very 
tip of a branchlet- Careful examination of a thoroughly moistened, 
relaxed branch apex under high magnification (about 30 to 40 X) 

usually reveals a few young leaves retaining the hairlike, white 

6. Selaginella lepidophylla (Hook. & Grev.) Spring in ]Mart. 

FLBras. 1(2): 126. 1840. 

Lycopodium lepidophyllum Hook. & Grev. Bot. Misc. 3: 106. 1832. 

On limestone ledges, on talus slopes, in crevices in both cal- 
careous and igneous rocks, on north-facing slopes and shaded sides 
of boulders at contact between gravelly soil and the rock, from 
western Texas and New Mexico south through ]\Iexico to El 

Specimens EfXAMiNEO: Baja California: Peninsular divide W of Caduano, 
Sierra de la Victoria, Cape Region, alt ca 4000 ft, 11 May 1959, Thomas 7859 
(DS) ; On north-facing cliff overlooking river at Las Animas Ranch, near base 
of El Picacho, Sierra de la Laguna, alt ca 2000 ft, 12 Oct 1941, Hammerly 281 
(CAS, DS). 

The plants of both this and the following species often are 
called "Resurrection Plant^* owing to the rapidity with which they 
flatten out and become bright green after being soaked by rain, 
in striking contrast to the apparently dead, compactly incurled 
branches during a period of drought. They can survive for months 
in the dormant condition, with the whole plant rolled into a 
grayish or brownish ball. The plants of both species are attached 
to the soil or rock only at the center of the rosette; no rhizophores 

160 American Feen Journal 

appear along the under side of the flattened branches during 
periods of active growth and photosynthesis. Dead leaves and 
branchlets from a plant itself, vegetative debris from adjacent 
plants, dust, and soil accumulate around the bases and within the 
curled up balls of dry ones. This detritus often complicates the task 
of making acceptable herbarium specimens. 

7. Selaginella pallescens (Presl) Spring in Mart. Fl. Bras. 

1(2): 132. 1840. 

Lycopodmm pallescens Presl, Rel. Haenk. 1: 79. 1825. 

Selaginella cnspidata (Link) Link, Fil. Sp. Hort. Reg. Bot. Berol. Cultae 
158. 1841. 

Around rocks, at bases of cliffs and rocky ledges, on gravelly 
soil of meadows and in open stands of shrubs, Sonora and Chi- 
huahua, southward in INIexico and Central America into northern 
South America, 

Specimi>ns examined: Baja California: Canyon at La Laguna, Sierra de la 
Lagima, E of Todos Santos, alt 1650 m, 25 Dec 1947, Carter, Alexander A 
Kellogg 23S5 (DS) ; Shaded rock faces along trail to Laguna from W, especially 
near summit, Sierra de la Victoria, 20 Aug 1955, Chambers 935 (DS); Potrero 
de Almerta, Arroyo de Almerta, E slopes of Sierra de la Victoria, inland from 
Caduano, among rocks and palms, alt ca 3400 ft, 9-1 1 May 1959, Thomas 7816 
(DS); Large meadow surrounded by oaks and Pinus cembroides, La Laguna, 
Sierra de la Laguna, alt ca 6000 ft, 16-18 May 1959, Thomas 7930C (DS). 

Selaginella pallescens is aptly named, for when curled up in the 
dry condition, the under sides of the branches and the leaves on 
that side are silvery white, whereas the compact ball of a dry plant 
of S. lepidophylla has a reddish brown to rusty tinge. In both 
species the leaves on the under side of the branches are consider- 
ably longer that those on the upper side, the lower leaves extending 
appreciably beyond the outer margins of the upper ones. The 
stoutish, denticulate-scabrous seta at the apex of each leaf in 
S, pallesceris contrasts sharply with the merely acute leaf of S. 
lepidophylla, which lacks a seta. This character can be seen easily 
with a hand lens, or in good light with the naked eye. Thus the 
two are not at all likely to be confused with one another. 

Dudley Herbakium, Stanford University, Stanford, CA 

Ferns of Dominica and St. Vincent IGl 

Notes on the Ferns of Eominica ar.d St. Vincent, IF 

C, V. Morton and David B. Lellinger 

This paper extends and concludes our notes on Lesser Autilleaii 
ferns and is based principally on collections made under the 
Bredin-Archbold-Smithsonian Biological Survey of Dominica that 
was active from 1963 to 1969, and also on collections made by 
C. V. Morton on St. Vincent in 1947. Since the prior report (Mor- 
ton & LelHnger, 1967) a few more botanists have visited Dominica 
and further researches by ourselves and others have solved some 
additional problems of taxonomy and nomenclature. 

DiPLAziUAi ambiguum Raddi, Opusc. Sci. Bologna 3: 292. 1819. 

Diplazium radicans sensu Christ, Farnkr. 220. 1896, pro parte. 

This South American species is known to us in the Antilles only 
from St, Vincent, where it was collected in the valley of the north 
fork of the Cumberland River at 400-600 m altitude {Morton 
5469) and in the upper valley of the Richmond River at 330-540 m 
altitude {Morton 6182). These specimens match an isotype of the 
species found ''in sylvis opacis prope Mandiocam, Brasiliae/' 
Raddi (FI, Morton photos 15914-15919 US). Diplazium amU- 
guum closely resembles D. striatum (L.) Presl, which is widely 
distributed in the Antilles. The former species is bipinnate- 
pinnatifidy at least in most of the lower pinnae, whereas the latter 
is strictly pinnate-pinnatifid. It also resembles D. expansum 
Willd., but lacks the usually abundant hairs on the lower surface 
of the costae and costules of that species and has entire, not 

ciliate-erose, indusia. 

Diplazium radicans (Swartz) Desv. is not a synonym of D. 
amhiguumy but is a name based on Asplenium radicans Swartz, 
1806, non L., 1759, (The two other names cited by Desvaux on p. 
281 of his 'Trodromus'' are invalid, later homonyms in Asplenium) 
Under A, radicans Swartz, Desvaux cites t. 76 of Schkuhr's 
''Kryptogamische Gewachse,'' a dubious species according to 
Christensen's ''Index FiUcum," plus Swartz' original publication 

* Partial cost of publication borne by the Smithsonian Institution. 

162 American Fern Journal 

of the species, which is based on one of his own specimens from 
Jamaica. Judging by Maxon's photograph of this probable holo- 
type and his annotation of the specimen, it is a taxonomic synonym 
of Asplenium dayi Hieron., which is the vahd name for this ex- 
tremely rare Jamaican plant. 

Elaphoglossum pusillu-m (Mett. ex Kuhn) C. Chr., Ind. Fil. 
314. 1905. 

Tliis species has heretofore been known only from the Greater 
Antilles and from Guadeloupe and Martinique. The first record 
for Dominica {Wilbur 8097) is from the forested slopes just below 
the summit and from dense thickets on the summit of Morne Trois 
Pitons, at 4000 4400 ft altitude. It is recorded as a common 


Gray Herb. 200: 115. 1970. 
Chambers 2596, from the summit of :Morne Trois Pitons, is the 
first collection of Eriosorus from Dominica. It was collected in a 
habitat typical for this genus: in a dense growth of pendent moss, 
on the side of a large, exposed rock at the lower margin of the most 
dwarfed pigmy forest, at about 4400 ft altitude. tartarea (Cav.) :\Iaxon, Contr. U. S. Nat. Herb. 

17: 173. 1913. 

Proctor (1965, pp. 219-220) has argued that the correct name 
for this species is P. ebenea (L.) Proctor, based on Acrostichum 
ebeneum L. Sp. PI. 1071. 1753. He lectotypifies this name on the 
basis of sheet 1245.14 in the Linnaean Herbarium, which represents 
a small, depauperate specimen of P. tartarea. We disagree, and 
consider .4. ebeneum to be a synonym of A. calomelanos [= Pity- 
rogramma calomelanos (L.) Link]. This has been the usual dis- 
position of A, ebeneum. 

The name A. ebeneum originated in a pre-Linnaean dissertation 
defended by J. B. Heiligtag in Linnaeus' "Amoenitates Academi- 
cae." It included several phrase-name synonyms based on material 
from Jamaica, Martinique, and Brazil, plus a description of a 

Ferns of Dominica and St. Vincent 163 

large specimen apparently agreeing with P, calomelanos (''Stipes 
pedalis k bipedalis'') that Linnaeus omitted from the account of 
A. ebeneum in the '*Species Plantarum." In that work the proto- 
logue is restricted to the following elements: 

1. A citation (Amoen. Acad. 1: 272. 1749) with the phrase 
name ''Acrostichum fronde pinnata, pinnis sessilibus oblongis 
sinuatis, summis brevissimis integerrimis" copied exactly from the 
''Amoenitates.'' Of the phrase-name synonyms in the '^Amoeni- 
tates," all but the two citations of Sloane's names, which are the 
only ones based on Jamaican material, should be disregarded for 
reasons set forth below in point 4. Sloane's names, descriptions, 
and plate are referable to P. calomelanos. The plate depicts a small, 
rather depauperate specimen. 

On the other hand, those Sloane descriptions and plate that 
truly do refer to P. tartarea (Voy, Jam. 35, t. 7,f. 1, 1707 and Cat. 
Jam. 21. 1696) are not mentioned by Linnaeus either in the 
^'Amoenitates^^ or in the "Species Plantarum/' 

2. A direct reference to Sloane, Cat. Jam. 20. 1696 and to Voy. 
Jam. 92, t, 53, f, 1. 1707, which are the two Sloane references in 
the '^Amoenitates'' mentioned above in point 1. 

3. A direct reference to Ray, Hist. PL Suppl. 83. 1704, which 
is based on Sloane, Cat. Jam. 20. 1696, and thus also refers to P. 

4. ''Habitat in Jamaicae sepibus humidiusculis," a statement 
of locality that restricts type material — specimens or literature — 

to Jamaica. The specimen in the Linnaean herbarium (1245.14) 
may or may not be a part of the protologue. Although it is anno- 
tated ''Aerosticum [sic] 20. ebeneum'^ in the hand of Linnaeus, 
there is no evidence that it is from Jamaica or was in Linnaeus' 
possession when either the ''Amoenitates'' or the ^'Species Planta- 
rum'^ was written. 

The phrase name in the ''Amoenitates" that Linnaeus copied 
in the ''Species Plantarum" does seem to apply to P. tartarea 
better than to P. calomelanos (''pinnis sessilibus oblongis sinuatis"), 
and it could apply to the specimen in the Linnaean herbarium. 
But the more ample description in the "Amoenitates" that Lin- 

164 American Fern Journal 

naeus omitted in the ''Species Plantarum" dearly does not refer 
to the specimen now in the Linnaean herbarium. The fact of this 
omission and the restriction of the type to Jamaican material 
indicates that Linnaeus' concept is based primarily on t, 53, f. 1 

of Sloane's ''Voyage." 

Tryon (1962, p. 60) suggested that Jenman (1886, p. 3S) in- 
ferentially lectotypified A. eheneum on the Sloane plate. However, 
Jenman's paper is merely an attempt to id(^ntify the Jamaican 
ferns in Sloane's herbarium; thc^re are no synonymies in taxonomic 
form. Under Gymnogramme calomelanos Jenman places the Sloane 
phrase name and reference (Hist. Jam. 92, t, 53, /. 1. 1707), and 
adds "Acrostichum ebeneum Linn. — a young plant [of G. calo- 
vwlanos],^^ This, we believe, was Linnaeus' actual concept. 

Pteris arborea L. Sp. PI. 1073. 1753. 

Hodge (1954, p. 85) lists this species only for the islands adja- 
cent to Dominica, but it has been collected several times on the 
island at 1000-1800 m altitude {Hodge 3236 from near Salybia, 
Chambers 2535 and LelUnger 393 from near Pont Cass^, LeUinger 
443 from near Trafalgar Falls, and Hodge 74 from Lisdara) . 

The pedate and areolate species of West Indian Pteris related to 
P. altissima Poir. have long been confused, and many names have 
been in use for them. Specimens in the U. S. National Herbarium 
can be separated easily with the following key: 

Fronds mostly 0.5-1 m long; costular areolae 2 (or 3 in large specimens), 
always parallel to the costa; costular spinules pointed and usually spread- 
ing; laminae gray-green P. altissima Poir. 

Fronds mostly 1-2.5 m long; costular areolae 1 or 2, the distal one, if present, 
always parallel to the costule, not the costa; costular spinules absent, 
rudimentary, or weakly developed and usually parallel to the costae; 
laminae bright green P. arborea L. 

For the most part, we have seen type fragments, authentic 
specimens, or sometimes photographs of type specimens of the 
names we place in synonymy. Under P. altissima Poir. we include 
P. brevinervis (Fee) Jenm., P. bulhifera Jenm., P. elata Agardh, and 
P. kunzeana Agardh. Under P. arborea L. we include P. aculeata 
Swartz, P. crassipes Agardh, P. laduca Poir. in Lam., P. longi- 
brachiata Agardh, P. multiserialis Jenm., and P. regia Jenm, 

Ferns of Dominica and St. Vincent 165 

The basis of P, arhorea should be t. 5 of Phimier's ''Traetatus do 
FiUcibus Americaiiis." (In this same work t. 11 is sterile material 
of the same species, but was not cited by Linnaeus.) The other 
basic reference cited by Linnaeus, /. 3 of Plumier's "Description 
de Plantes de TAmerique/' is also part of the protologue of 
Polypodium spinosum L., presumablj^ through a typographical 


Pteris hartiana Jenm. is a taxonomically distinct species from 
Trinidad. It has at least three areolae along the costae, one long 
and the others shorter; as in P, althsima, all are parallel to the 
costa. It differs from P. altissima in having 4-6, rather than 2 or 3, 
rows of areolae between the costae (or costules) and the margin. 

Literature Cited 

Hodge, W. H. 1954. Flora of Dominica, B. W. I. Part. I. Lloydia 17: 1-238- 
Jenm AN, G. S. 1886. On the Jamaica ferns of Sloane's herbarium. J. Bot. Brit. 

& For. 24 : 33-43. 
Morton, C. V. and D. B. Lellinger. 1967. Notes on the ferns of Dominica 

and St. Vincent. Amer. Fern J. 57: 66-77. 
Proctor, G. R. 1965. Taxonomic notes on Jamaican ferns. Brit. Fern Gaz. 

Tryon, R. M., Jr. 1962. Taxonomic fern notes. 11. Contr. Gray Herb. 185: 


U. S. National Herbarium, Smithsonian Institution, 
Washington, DC 20560. 

166 American Fern Journal 

Two New Tree Ferns from Costa Rica 

Luis Diego Gomez 

The tree fern flora of Costa Rica is richly represented by some 
50 or 55 species of the Cyatheaceae and Dicksoniaceae. Two 
have been knowTi from Cocos Island (o°30'N, 87°03'W), a Costa 
Rican possession in the Pacific Ocean some 300 miles southwest 
of the Punta Burica Peninsula, the southernmost part of Costa 
Rica. These species, Cyathea notabilis Domin and Trichipteris 
7iesiotica (Maxon) Tryon (formerly known as Alsophila notabilis 
Maxon and A. nesiotica Maxon), occur on the island from sea- 
level to its highest point, Monte Yglesias, about 900 m altitude. 
During an eight-week survey of the island's cryptogamic flora, I 
noticed a tree fern that differed in several respects from the two 
previously reported species. Further study lead to the conclusion 
that this is a new species: 

^Oyathea alfonsiana Gomez, sp. nov. 

Truncus usque ad 1.5 m altus, ca 5 dm diam, apice ut in petio- 
lorum basibus dense paleaceus, paleis oblongo-lanceolatis, bi- 
coloribus, medio rubescenti-avellaneis, marginibus cremaei-albis; 
foliorum cicatrices obviae; stipites interdum pcrsistentes, supra 
ut in rhachi canaliculati et pilosi, subtus glabri, spinulis sparsis 
parvis_ apice fere rotundis; pneumatophora non obvia; costae 
primariae- supra pilosae, subtus glabrae, secundariae supra pilosae, 
subtus paleaceae, paleis numerosis parvis appressis citrini-hyalinis 
membraneis bulliformibus; pili petiolorum supra rhachium costa- 
rum costularumciue multicellulares fcrruginei attenuati crassi 
rigidi apicem versus curvati; laminae foliorum oblongo-lanccolatae, 
1.4 m longae, 80 90 cm latae, bipinnato-pinnatifidac; pinnae 
basales abrupte reductae, apice pinnatisectae; pinnae 12-15-jugae 
alternae, 35-40 cm longae, 12-15 cm latae, pinnulis usque 21- 
jugis altcrnis vel subalternis, 6-8 cm longis, 2 cm latis, fere ad 
costam sectis, segmentis usque 14 paribus alt(;rnis oblongis, basi 
paullo constrictis, 7-9 mm longis, 4-5 mm latis, lacviter lobatis, 
apice subcrenulatis, venis 7- vel 8-jugis, 1-3-plo furcatis; sori 
mtramediales, indusiati. Indusium squamiforme, parvum. 

•^YPE: Twin Arnnntainc. TTr^r^^^ 



American Fern Journal 

Volume 61. Plate 25 

J -> 







_ L 



1 -■ 

- V 



HoLOTYPE OF Cyathea alfonslvna (CRj. FiG. 1. Medlvl pinnae. Fig. 2 
Medl\l pinnules. Scale in cm. 

168 American Fern Journal 

Paratypes (identity confirmed by R. M. Tryon, Jr.): Cocos 
Island, Holdridge 5160 (GH), W, L. Schmitt 129, 130, 131 (all US). 

According to the recent classification of the Cyatheaceae by 
Tryon (1970), this species falls in Cyathea, mainly on the structure 
of the scales of the trunk, stipe, and rachis; that is, the scales are 
marginate with a broad margin of cells unlike those of the central 
portion. The stipe scales lack setae and have, like the rest, a 
filamentous apex. Various types of indument are present on the 
stipes, rachises, costae, and costules. The venation is free and there 
is a scale-like indusium present. The new species differs from (7. 
notabilis and from T. nesiotica in general habit, but especially in 
having a scale-like indusium; the indusium is absent in Trichipteris 
and is sphaeropteroid in C, notabilis. Such peculiarities of the in- 
dusia of these Cocos Island tree ferns were first noted by Maxon 
in the specimens collected by Schmitt (C. V. Morton, pers. comm.) . 
Cyathea alfonsiana is named after ]Mr. Alfonso Jimenez ^L, 
former Curator of the Costa Rican National Herbarium, as a 
humble tribute to his many contributions to the general knowledge 
of his country's flora. 

The second new species is an endemic fern of the central north- 
eastern slopes of the Talamanca Ridge in continental Costa Rica. 
This species is in evident danger of extinction due to uncontrolled 
deforestation. It presents a number of interesting characters, 
one of them of ecological interest. In collecting some immature 
sori for chromosome counts, I discovered that a moth of the 
Nymphalid group deposits its eggs in rows that duplicate exactly 
the disposition and appearance of the immature sori. All attempts 
to rear the larvae and adults of this remarkable insect failed, and 
the moth still remains unidentified. The new species is a conspic- 
uous, tri pinnate tree fern also belonging to Cyathea in Tryon' s 
classification. It was formerly described, but unfortunately not 
validly or efTectively published, by Nisman (1965) as Hemitelia 
holdridgei, and is here published as: 

Cyathea holdridgeana Nisman & Gomez, sp. nov. 

Truncus rectus ca 2 m longus, inermis, paleaceus, paleis stram- 
ineis ovato-lanceolatis, semiuncialibus, brunneis guttatis. Frons 

American Fern Journal 

Volume 61, Plate 26 


^ ^ 


Fig. 4. Medial pinnule and part of rachis. Scale in cm. 

170 Ameeican Fern Journal 

2 m longa, pinnatisecta. Stipes et rhachis semiteretes canaliculati, 
ochracei opaci, parce muricati. Stipes 7 cm longus, 1.5 cm crassus, 
ad basim paleaceus, primo furfuraceus, postea glabrescens; rhachis 
supra strigosa, subtus furfuracea, paucis paleis sparsis^ pinnis 17- 
jugatis. Pinnae petiolatae (petiolo 2.5 cm longo), rhachibus fere 
teretibus canaliculatis^ dense strigosis, subtus furfuraceis squamu- 
losiS; deltoideo-lanceolatis; pinnuHs 20-j ugatis ; laminae apice 
acuminatae serratae. Pinnulae petiolulatae (4 mm) dehoideo- 
lanceolatae, segmentis 14-jugatis. Costa dense strigosa; trichomata 
septata fulva, subtus furfuraceo-squamulosa. Segmenta in basali 
parte pinnulae distincta, sessilia, ad basim laeve inaequalia. Pars 
apicalis pinnulae pinnatisecta fere ad cost am. Laminae coriaceae, 
supra glabrae, virides, subtus pallidiores, margine revoluto serrato. 
Costa secundaria ad basim strigosa, ad apicem cum 2-4 pilis 
rigidis, subtus squamulosa. Sori inframediales, orbiculares, trans- 
lucenti-virides, costis secundariis propinqui. 

Type: La Chonta, at Ean 55 of the Interamerican Highway, 

Prov. Cartago, Costa Rica, alt 2200 m, Carmen Nisman S. 104 

(CR;isotypesF, GH). 

Paratypes: La Chonta, Prov. San Jose, Costa Rica, alt 2300 m, 
Gomez 25^2, 2577, 2560, Goldgeivicht & Gomez 2675 (all Herb. 
Gomez); Km 54 of the Interamerican Highway, Costa Rica, 
J. A. Saenz (Herb. Gomez). 

Although there are a number of species of Cyathea wdth tri- 
pinnate or tripinnate-pinnatifid fronds, such as C. hemiotis Christ 
and C hastulata Christ, the closer affinity of C. holdridgeana seems 

ultiflora J. E, Smith 


Kuhn) Domin, from which it differs in having sessile pinnae, 
petiolulate pinnules, a strictly inframedial sorus, and bicolorous 
and corticinate scales on the stipe. 

Literature Cited 

Nisman S., Carmen. 1965. Estudio taxonomico y ecologico de los helechos 

arborescentes (Cyatheaceae y Dicksoniaceae) de Costa Rica. Lie. 
Thesis, University of Costa Rica. 

Tryon, R. M., Jr. 1970. The classification of the Cyatheaceae. Contr. Gray 

Herb. 200 : 3-53. 

:Museo Nacional de Costa Rica, Apartado 749, San Jose, 
Costa Rica. 

Spore Germination and Prothallial Growth 171 

Comparative Habitat Requirements 

for Spore Germination and Prothallial Growth 

of Three Ferns in Southeastern Michigan 


Investigation of the physiological ecology of various plants has 
yielded information concerning mechanisms and patterns of 
evolutionary divergence (Bjorkman & Holmgren, 1963; Ayodele 
Cole, 1967; Hadley & Levin, 1969; Mooney & Billings, 1961). 
Such studies have largely omitted pteridophytes. 0\\ing to the 
alternatioii-of-generations life cycle, certain problems arise con- 
cerning physiological divergence within groups of pteridophytes. 
The success of pteridophytes in a community is obviously limited 
both by the successful establishment of free-living gametophytes 
and the survival of sporophytes. Thus, the habitat requirements 
for spore germination and prothallial growth are critical factors 
for the success of the plant. Physiological divergence between 
populations or species, as a result of selective pressures of eco- 
logically different habitats, should be apparent in both phases of 
the pteridophyte life cycle. 

Farrar (1967) found fern gametophytes reproducing inde- 
pendently in the southern Appalachians, far north of the normal 
ranges of their sporophytes. Conway (1953) stated that there are 
few records of finding sporelings of Pteridium aquilinum in Scot- 
land, even though the sporophytes are common and spore pro- 
duction is heavy. She suggested that spores are released late 
during the growing season and that Pteridium sporelings are more 
susceptible to frost than are those of other species observed. 
Conway also noted that sporehngs are subject to attack by soil 
insects and fungi. These two studies suggest that the gameto- 
phytic and sporophytic phases of a plant can have different 
habitat requirements and that one phase of the Hfe cycle can be 

^ I wish to acknowledge the guidance of Dr. Warren H. Wagner, Jr. and 
Dr. Edward L. McWilliams and the use of the facihties of the Matthaex 
Botanical Gardens, University of Michigan. 

172 Amekican Fern Journal 

limiting to the success of the species in extending its geographical 
range and ecological amplitude. 

The purpose of this investigation is to compare the habitat 
requirements of prothallia of three polypodiaccous species that 
occupy ecologically contrasting habitats in southeastern JNIichigan. 


Species from habitats differing in exposure to sunlight and in 
substrate pH were selected for study. The habitats are located in 
Washtenaw County, Michigan. Thelypteris palustris Schott 
(Marsh Fern) grows abundantly on the open sphagnum mat of 
Mudlake Bog. Woodwardia virginica (L,) J. E. Sm. (Virginia Chain 
Fern) occupies floating, graminoid mats in a swamp clearing at the 
Waterloo Recreation Area. Adiantum pedatuvi L. (Maidenhair 
Fern) grows in an upland northern woods at Sharon Hollow. 

Light intensity and temperatures were taken at the habitats. 
Light intensity was measured \\dth a General Electric Type DP-9 
light meter and temperatures were taken at 30 cm and at the sub- 
strate surface with glass mercury thermometers. Readings were 
taken at each habitat and at an adjacent, gravel road to obtain 
habitat and ambient conditions, respectively. The ambient 
readings were intended to facilitate comparisons of environmental 
data taken on different days, August 7 at Sharon Hollow and at 
Waterloo and on August 30 at Mudlake Bog. Both days were 
bright and hot, as indicated by the similarity of the ambient 
conditions of the habitats. These data appear in Table I. 

Spores were collected by washing fertile fronds under running 
tap water, blotting them dry, and placing them in envelopes for 
storage for several months at 6"" C. Upon removing the spores from 
storage, they were filtered through a 0.177 mm sieve and several 
layers of cheesecloth to screen out extraneous material such as 
sporangia. They were then sown in 10 X 35 mm plastic petri 
dishes containing nutrient medium^ at a density of approximately 
100 spores per square cm. The medium was one of several varia- 
tions of full-strength Bold's nutrient solution. Liquid medium and 
liquid solution added to blotting paper were usable, but these 

Spore Germination and Prothallial Growth 173 

techniques often led to fungal or algal contamination of the cul- 
tures. Bold's solution solidified with 1 % bacto-agar was the 
medium least susceptible to contamination. 

The petri dishes Avere placed under gradients of light intensity 
and pUj inside a growth chamber^ and of temperature on a thermo- 
gradient plate. In the light intensity experiments, the light source 
was a combination of incandescent and Cool White fluorescent 
lamps; the petri dishes were placed at varying distances from the 
light source and/or were covered with layers of cheesecloth in order 
to vary the light intensity. Temperatures inside the plates were 
determined with a Telethermometer probe and maintained at 
25 d= l^'C. Temperature control at the higher values of light 
intensity was facilitated by placing the dishes in the direct flow 
of cool air above the chamber vents and by positioning a plexiglass 
water screen containing 10 cm of Avater between the dishes and the 
light source. The pH of the medium was adjusted to 6.7 by adding 
NaOH or HCl for both the light and temperature experiments. 
Petri dishes were placed under 500 ft-c at 25 ± 1°C in the pH 
experiments and under incandescent Hght at 500 ft-c in the tem- 
perature experiments. 

After a growth period of 3 weeks, the percentage of germination 
and prothalHal width (to 0.1 mm) were determined, the former 
based upon 400 spores or prothaUia and the latter upon the maxi- 
mum width of 40 prothaUia, measured with an ocular micrometer 
mounted in a dissecting microscope at 40 X. ProthalHal width 
was chosen as an index of growth because width increments of 
gametophytes appeared to be more directly related to increments 
of prothallial area than were length increments. Dark-grown 
prothallia in the light intensity experiments, for example, were 
long and spindly, whereas cordate prothallia at higher light in- 
tensities attained somewhat greater prothalHal width but only 
sHghtly greater length than dark-grown prothallia. 

Environmental Data 

Table I shows that hght intensity (LI) was highest in the open 
habitats. The bog habitat is fully exposed to hght except for 


American Fern Journal 

occasional shade cast by poison sumac shrubs; the swamp habitat 
is shaded only during the early morning hours by an upland oak- 
hickory woods just east of the swamp. LI at the \vooded habitat 
on August 7 varied from 77 ft-c in deep shade to 1192 ft-c where 
light flecks penetrated the canopy, even though ambient LI (at 


the road) was 5700 ft-c. Table I also shows that the air and surface 
temperatures, relative to ambient temperatures, Avere also higher 
in the open habitats than in the wooded habitat. 

Table I shows that the open habitats are acidic, whereas the 
woodland habitat is circumneutral. The bog — a typical northern 

Table I. Comparison of the Habitats of Thrbe Fern Species 




Light Intensity (ft-c) 


Temperature ("C) 

ambient (30 cm) 

30 cm 
Substrate pH 

Sharon Hollow 
wooded upland 





rich humus 

Mudlake Bog 
open bog 






swamp clearing 




sphagnum mat graminoid mats 



one— is dominated by sphagnum, larch, and ericaccous shrubs. 
The swamp substrate is acidic and highly anaerobic. The wood- 
land substrate consisted of leaf litter overlying a deciduous forest 


Thelypteris and Woodwardia spores required about 2 days for 
germination, whereas Adiantum spores germinated in 5 days. The 


first two species also had faster growth rates than did Adiaritwi 
during the 3-week period following germination. Prothalhal width 

Spore Germination and Prothallial Groavth 175 

of Adiantum never exceeded 0.6 mm, whereas that of the other 
species reached 2,0 mm at the lower pH values. The shorter 
germination time and faster growth rates of the open-habitat 
species may constitute an adaptive advantage. It is likely that 
prothalha of these species would have faster growth rates and be 
more competitive than would Adiantum m an exposed habitat. 

Temperature Experiments 

Petri dishes of each species were placed on a thermogradicnt 
plate to establish a IS-So'^C temperature gradient. In addition, 
dishes were also placed in two cold rooms at 5 and 10°C. As seen 
in Fig. 1, none of the spores germinated at 5°C, although spores 
did germinate after they were moved to 25°C following their 
exposure to the cooler temperature. Other experiments showed 
that none of the spores germinated at 40°C and that the spores lost 
viability, for they did not germinate when they were moved to 
25°C. Thelypteris and Woodwardia spores germinated at 35°C, 

but Adiantum spores did not. 

The temperature hmits of germination for Thelypteris and 
Woodwardia are approximately 10-35°C, whereas those of Adian- 
turn are about 10-30°C. The higher temperature tolerance of the 
open-habitat species may be considered adaptive, since surface 
temperatures of the bog and swamp probably reach 30-35*'C 
regularly during the mid-hours of the day. Surface temperatures 
of the woodland habitat probably seldom reach 30°C. Spores 
landing on the sphagnum mat of the bog or on the floating mats of 
the swamp would certainly be exposed to more adverse temper- 
atures than those landing on the forest floor of the woodland 


Figure 1 also shows that optimum or near-optimum tempera- 
tures for prothallial growth are higher for the open-habitat species 
than for Adiantum. These were 25-30^C for Thelypteris, 25°C for 
Woodivardia, and 15-25°C for Adiantum. Prothallial growth of 
the three species followed much the same pattern as germination, 
although it can be seen that the temperature hmits of germina- 
tion are broader than those of maximum or near-maximum growth 
of prothalha. Although spores of Thelypteris and Woodicardia 


American Fern Journal 

germinated at 10°C, prothallial growth was relatively low at 
temperatures below^ 25°C. By contrast, Adiantum prothallia had 
relatively high growth rates at 15*^0. The preference for lower 
temperatures in the latter species, as compared to the open- 
habitat specieSj appears to be an adaptation to the cooler wood- 
land habitat. Growth of Adiantum prothallia, compared to the 
other species, w^as decidedly slower and dropped abruptly at 
temperatures above 25°C. The preference for higher temperatures 








10 15 20 25 30 

Temparatura (C ) 

♦ maan 

• 1 standard deviation 

1 ranga 

Fig. 1. The effect of temperature on prothallial width and percent 
germination of spores. 

and the tolerance of higher temperatures in Thelypteris and 
Woodwardia appear adaptive to the open habitats of these species. 
Thus the temperature requirements for prothaUial growth of these 
species are even more in accord Avith their respective habitat condi- 
tions than are their temperature requirements for germination. 

The temperature limits of spore germination may be compared 
with those reported by Hartt (1925) for Onoclea sensibilisj in 
which the spores germinated at 16-34°C with an optimum of 28''C. 
Spore viability was lost at 40°C. Hevly (1963), in a study of the 
physiological ecology of cheilanthoid ferns, reported much higher 

Spore Germination and Prothallial Growth 177 

temperature limits. Spores of Notholaena cockisensis, a xerophytic 
fern that typically grows fully exposed to the sun, germinated at 
10-50°C with an optimum of 25-35°C, whereas Cheilanthes fend- 
lerij which usually inhabits crevices around the bases of boulders, 
germinated at 0-35°C with an optimum of 15-20°C. 

The spores of the species investigated here germinate in only a 
few^ days after sowing, and previous experiments showed that 
freshly collected spores are fully capable of germinating. These 
observations suggest that spores would germinate in the field soon 
after their discharge from sporangia, and that the overwintering 
stage is the prothallium rather than the spore. Observations of 
seasonal phenology of these species showed that initial dehiscence 
of sporangia occurred on June 22 for Woodwardia, August 7 for 
Adiantum, and August 25 for Thelypteris. 

It would seem reasonable that spores shed late in the gro\ving 
season, such as those of Adiantum and ThelypteriSj w^ould over- 
winter before germinating. If this were the case, the spores would 
probably have a chill requirement for germination. This was not 
found to be the case for these species, but the question of w^hether 
spores that fail to germinate during the growing season in which 
they were discharged could survive winter conditions arises. It 
was noted that the spores retained their viabiUty after several 
weeks of storage at 6°C and at shorter exposure to 5°C on agar 
medium. In an attempt to resolve this question, spores were 
sow^n in a liquid medium which was then frozen for 30 days and 
then brought to room temperature. A high percentage of the spores 
germinated, and in the time periods characteristic for each species. 
It appears that spores of these species are able to retain viability 
under winter conditions, although the evidence suggests that the 
usual overwintering stage is the gametophyte rather than the 
dormant spore. These are only tentative observations and need 
to be followed by intensive field studies. 

Light Intensity Experiments 

A LI gradient of 100-3000 ft-c w^as established. In addition, 
petri dishes were covered with black cloth, after receiving an 


American Fern Journal 

initial light stimulus during sowing, to effect dark conditions. 
Germination percentages and prothallial growth for the species 
over the gradient are shown in Fig, 2, 

Although spores of all species germinated at all LI values, the 
percentage of germination w^as lower in darkness and high germi- 
nation rates were reached at 100-3000 ft-c for all three species. 





Light Intensity ( " 10^, f-c ) 

• mean 

• 1 standard deviation 

X range 

• ^—germination 

Fig. 2. The effect of light intensity on prothallial width and per- 

The prothallia of all three species were long and spindly when 
grown in darkness. The prothaUial width of Thelypteris was 
greatest at 1000 ft-c, and that of Woodwardia and Adiantum at 
500 ft-c, but prothallial growth dropped off more abruptly at 
values above 500 ft-c in Adiantum than in the open-habitat species. 
The relatively faster growih rates of the open-habitat species, as 
compared to those of Adiantum^ suggest their selective advantage 
in establishment in open communities. 

The apparent chlorophyll levels of prothallia also indicate that 
the open-habitat species have higher tolerance to high LI than 
does Adiantum, Spores of all three species germinated at 3000 ft-c, 

Spore Ger:\iination and Prothallial Growth 179 

but the resulting prothallia were chlorotic as well as small. At 
2000 ft-c, prothallia of Thelypteris and Woodicanlia were pale 
green, whereas those of Adiantuni were devoid of green color. 
Darker green colors were found in all three species at 100-500 ft-c. 

Spores of all three species germinated in darkness after an 
exposure of only a few seconds to the fluorescent light used during 
sowing. Mohr (1963) stated that Hght is an almost universal re- 
quirement of spore germination and prothalUal development 
among ferns. Borth\\ick et al. (1952) noted that certain small 
seeds require a dose of red light for germination, a stimulus that 
can be nullified by a subsequent treatment with a far-red light. To 
determine whether spores of the species under study have no 
light requirement for germination, or whether the brief exposure 
of the spores during sowing to fluorescent hght (including red 
hght) satisfied a possible hght requirement, spores were soato under 
neutral hght only (absence of red hght) and then placed in darkness 
After a period of two weeks, no spores had germinated, indicating 
a red-light requirement of these spores. It is hkely that the germi- 
nation of these species is under operation of a red and far-red 
reversible photoreaction. 

Du Buy and Neurnbergk (1938) state that Pteridhm aquilinum 
spores germinate well in hght of ''high intensity." This is in accord 
with results reported here for Woodwardia and Thelypteris, but 
Hevly (1963) found that desert ferns of the Southwest germinated 
at a somewhat lower maximum LI. Miller and Miller (1961) 
found that prothallia of Onodea sensibilis grew much more rapidly 
under 400 ft-c than under 220 or 28 ft-c. It was also observed m 
the present study that prothalha of all three taxa grew faster 
at 500 ft-c than at lower LI values. 

pH Experiments 

Spores were placed in a pH gradient of 4-10 (Fig, 3). Thelypteris 


requirements than did Adiantum, whose woodland habitat is 
circumneutral. Spore germination of Thelypteris was high over 
the entire gradient, but prothallial growth showed a marked pref- 


Amekican Fern Journal 

erence for pH 5 and decreased sharply at higher pH values. 
Assuming that the germination process is adapted to the natural 
habitat of this species, this is in agreement with the observed pH 
value of 5.5 for water squeezed from sphagnum of Mudlake Bog. 
Woodwardiaj whose habitat substrate pH was determined to be 
5.9, had both the highest percentage of germination and the 


• 1 standard deviation 

J rang© 

■ %-germi nation 

Fig. 3. The effect of pH on prothallial width and percent germina- 
tion OF SPORES. 

fastest growth rates at pH 5-7. Percent -germination for Adiantumy 
whose soil pH was 7.6-7.8 in several samples, was high at all pH 
values except pH 4, and prothaUial growth was maximal at pH 8. 
Again these species differences appear to reflect habitat differences. 
Fig. 3 shows that maximum prothallial width of Thelypteris 
and Woodwardia exceeded 2.0 mm at low pH values, whereas 
prothallia of these species grown in the temperature and LI 
experiments were smaller. This is probably a result of the pH (6.7) 
used in the latter experiments, which is higher than the optimum 

Spore Germination and Prothallial Growth 181 

pH values for these species. The comparative growth rates of these 
three species are also very apparent in the pH experiments, as 
prothallia of Adiantum at optimum pH (8) reached a maximum of 
only 0.5 mm in diameter. This contrast dramatically demon- 
strates the faster inherent growth of the open-habitat species as 
compared to Adiantum. 

It has been accepted for some time that fern species have 
definite habitat requirements for substrate pH. \ATierry (1920) 
listed pH ranges for several rock ferns, and Kruckeberg (1964) 
reported that certain fern species of the Pacific Northwest are 
restricted to locaUzed ultramafic rocks. It is reasonable that 
gametophytes have pH requirements similar to those of the sporo- 
phytes. McMillan et al. (1968) attributed the narrow distribution 
of Pilularia americana in Texas partially to the inability of spores 
to germinate on limestone soils. Spore germination of this species 
was inhibited at pH values above 6.4, although sporelings trans- 
planted to limestone soils grew at normal rates. The habitat of 
Pilularia in central Texas has a pH of 5.2-6.1. 

Hevly (1963) found a correlation between the pH of habitats 
of certain cheilanthoid ferns and their corresponding pH re- 
quirements of germination. Notholaena cochisensis, which grows 
on limestone soils, germinates at pH 7-9 with an optimum of 
pH 8. Notholaena grayi germinates at pH 5-8 \\'ith an optimum of 
pH 6 and grows on soils on igneous rock. 

The different pH requirements of the open-habitat species and 
Adiayitum, like the different temperature and LI requirements of 
these species, reflects the different habitat conditions of these 
species. These observations provide strong evidence that the 
processes of spore germination and prothallial growth of these 
species, as a result of variation and selection, have undergone 
physiological divergence. 


Ayodele Cole, N. H. 1967. Comparative physiological ecology of the genus 

Eriogonum in the Santa Monica Mountains, Southern California. 
Ecol. Monogr. 37: 1-24. 

182 American Fern Journal 

Bjorkman, 0. and P. Holmgren. 1963. Adaptability of the photosynthetic 

apparatus to light intensity in ecotypes from exposed and shaded 

habitats. Physiol. Plant. 16: 889-914. 
BoRTHvvicK, H. A. ET AL. 1952. A reversible photoreaction controlling seed 

germination. Proc. Nat. Acad. Sci. 38: 662-666. 
Conway, Elsie. 1953. Spore and sporeling survival in Bracken (Pteridium 

aquilinum (L.) Kuhn). J. EcoL 41 : 289-294. 
Du Buy, H. G. and E. L. Neurnbergk. 1938. Growth, tropisms, and other 

movements, Chapter X in Manual of Pteridology, ed. F. Verdoorn. 

M. Nijhoff, The Hague. 

Farrar, D. R. 1967. Gametophytes of four tropical fern genera reproducing 

independently of their sporophytes in the southern Appalachians. 
Science 155: 1266-1267. 

Hadley, E. B. and D. A. Levin. 1969. Physiological evidence of hybridiza- 
tion and reticulate evolution in Phlox maculata. Amer. J. Bot. 56: 

Hartt, Constance E. 1925. Conditions for germination of spores of Onoclea 

sensibilis. Bot. Gaz. 79 : 427-440, pL XXIX. 
Hevly, R. H. 1963. Adaptations of cheilanthoid ferns to desert environments. 

J. Ariz. Acad. Sci. 2: 164-175. 
Kruckeberg, A. R. 1964. Ferns associated with ultramafic rocks in the 

Pacific Northwest. Amer. Fern J. 54: 113-126. 
McMillan, C. et al. 1968. Factors influencing the narrow restriction of 

Pilularia americana in Texas. Southw. Nat. 13: 117-127. 
Miller, J. H. and P. M. Miller. 1961. The effect of different light conditions 

and sucrose on the growth and development of the gametophyte of 

the fern, Onoclea sensibilis. Amer. J. Bot. 48: 154-159. 
MoHR, H. 1963. The influence of visible radiation on the germination of 

archegoniate spores and the growth of the fern protonema. J. Linn. 

Soc. London, Bot. 58 : 287-296. 
MooNEY, H. A. and W. D. Billings. 1961. Comparative physiological ecology 

of arctic and alpine populations of Oxyria digyna. EcoL Monogr. 

31 : 1-29. 

Wherry, E. T. 1920. The soil reactions of certain rock ferns. I, II. Amer. Fern 

J. 10: 15-22, 45-52. 

Department of Botany, University of ^Michigan, Ann 
Arbor, Ml 48104. 

Gametophyte of Thelypteris erubescens 1S3 

The Gametophyte of Thelypteiis erubesceis 

Lenette R. Atkinson erubescens, the Reddening Thelypteris, is a fern of 
the mountains with a thick, creeping rhizome and a leaf that can 
be 6 feet tall or more. The stipe and large veins may be purplish- 
tawny (Beddome, 1892, p. 289) or wine-red when dried (Ching, 
19G3, p. 320). It is one of a small group of ferns in North India and 
West China and has been known for some time, Iwatsuki (1965, 
p. 178) in his study of the thelypteroid ferns says of it, the "dis- 
tinctness of this large and handsome fern has been recognized by 
various authors since WalHch (1828) gave this name to his Nepalian 
specimen." Ching (1963, p. 320), who had examined the Wallichian 
collection (see Ching, 1936, p. 294), says 'Tt has been long noted 
that of all the Chinese thelypteroid ferns, the well-known ex- 
indusiate Dryopteris erubescens (Wall.) C. Chr. from the China- 
Himalayan region always appears to be foreign among all the 
known genera, in none of which can it possibly fit." Wallich called 
it Polypodium; other authors, depending on which characters were 
thought to be more important, have placed it in Glaphyropteris, 
Phegopterisy Nephrodium, and Dryopteris. Those who today would 
place this fern in Christensen's tropical American subgenus 
Glaphyropteris (on the basis of habit and aerophores present) 
would do so against Christensen's (1913, p. 158) judgement. In 
discussing his new subgenus Glaphyropteris he says "I do not know 
any Old- World's species which can safely be referred hereto. The 
Himalayan D. erubescens (Wall.) C- Chr. is similar in habit but 
otherwise quite different/' Ching (1963, p. 320) concurred and 
placed it in his newly created genus Glaphyropteridopsis, the type 
species of which is Thelypteris erubescens. The chromosome num- 
ber reported for this fern by Mitui (1968, Lasfrea erubescens) is 
n = 72 (on the base 36, one of the more common of the various 
numbers reported for the Thelypteridaceae). The gametophyte 
also shows the general characteristics of the group. 

I am indebted to Dr. R. E. Holttum for spores of Thelypteris 

184 American Fern Journal 

eruhescens (Wall, ex Hook.) Ching from a plant from Nepal growing 
at Kew Gardens, England. Dr. Holttum writes me that this 
"Kew plant . . . was much smaller than the size to which plants of 
eruhescens commonly attain." He continues, "In the Kew her- 
barium are several quite small specimens, much like that from 
which your spores came and also some very much larger, with 
pinnae to 35 cm. long and 3 cm. wide. There are also some rather 
intermediate in size.'^ The gametophytes were grown under fluores- 
cent light on Knop's solution solidified with agar (Atkinson, 1970). 

The bilateral spores are brown singly, black in mass and measure 
about 33 X 21.5 ju, exclusive of the spore coat. They are orna- 
mented by more or less discrete little wings which when dried are 
sharp-pointed and give the spore a shaggy or spiny appearance. A 
colorless rhizoid emerges first at germination. A 4-6-celled filament 
is formed whose cells may be greatly elongated when developed 
from spores retained and germinating within the sporangium. The 
first division initiating the cell plate is usually oblique in the 
terminal cell of the filament; occasionally it is vertical and in the 
subterminal cell. A cordate thallus is formed by the activity of an 
apical cell which may be replaced by an apical meristem as early as 
a month after germination. The margins of a month old thallus 
are wavy and may bear 0-3 hairs. At two months simple hairs 
may also be found on the surface. The thallus bears sex organs by 
3.5 months, has a broad cushion and large wings with irregular 
margins. At 6 months the wings are very large, with folds in the 
surface and shallowly notched margins. Regeneration in older 
thalli takes place from the margins of the wings, and little groups 
of cells project irregularly from the sides of the cushion on the 
ventral surface. These may bear glandular hairs and abortive 
antheridia, and are interpreted as regenerative outgrowths. 

The first hairs are simple, chlorophyllous and marginal. They 
are late in appearing; some cordate thalli at one month do not 
have any hairs. They become glandular and the secretion is yellow. 
At 2 months they appear on the surface near the notch but are 
not abundant in either position; one thallus with 2 surface hairs 
had none on the margin. 

Gametophyte of Thelypteris erubescens 185 

The rhizoids are colorless or palest tan, long but do not tangle, 
and are abundant on the cushion, hiding the older sex organs. 
When thalli overlap in culture, rhizoids are often borne over the 
lower part of the cushion on the dorsal surface. 

The sex organs are of the advanced type. Archegonia appear at 
2 months and embryos at 3.5 months, although no effort was 
made to produce them. Antheridia are scarce, and those first seen 
were on separate, usually smaller thalli. At C months they appear 
also at the sides of the cushion beside the opened and younger 
archegonia. Dehiscence was not observed. 

The large, green thallus with simple,, yellow glands, ruffled 
wings, abundant colorless or pale tan rhizoids, and sex organs of 
the advanced type is not out of place among the gametophytes of 
the Thelypteridaceae, but it may also be found elsewhere among 
the ferns. The spore is not very large (as is sometimes the case with 
polyploids), but, then, spore size at the species level may not 
always accompany an increase in chromosome number, as Bir 
(1966) has reported for some Himalayan ferns. Dr. Holttum sug- 
gests that a field study of Thelypteris enihescens is desirable in a 
place where the species grows abundantly. 

No notable gametophytic features are observed in Thelypteris 
enihescens to suggest relationships to other ferns Avithin the family. 

Literature Cited 

Atkinson, Lenette R. 1970. The gametophyte of Pleocnemia conjugata and 

of Arcypteri? irregularis. Phytomorphology 20: 78-83. 
Beddome, R. H. 1892. Handbook of the ferns of British India. Calcutta. 
Bir, S. S. 1966. Correhition between spore-size and polyploid-level in the 

Himalayan asplenioid and athyrioid ferns. J. Palynolog>^ 2: 41-48. 
Ching, R. C. 1936. A revision of the Chinese and Sikkim-Himalaya Dryopteris 

with reference to some species from neighboring regions. Bull. Fan 

Mem. Inst. Biol. 6: 237-347. 
. 1963. A reclassification of the family Thelypteridaceae. Acta 

Phytotax. Sin. 8: 289-335. 
Christensen, C. 1913. Monograph on the genus Dryopteris I. K. Danske 

Vidensk. Selsk. Skr. VII, 10: 55-282. 
Iwatsuki, K. 1965. Taxonomy of the thelypteroid ferns with special reference 

to the species of Japan and adjacent regions. IV. Enumeration of 

186 American Fern Journal 

the species of Japan and adjacent regions. Mem. Coll. Sci. Univ. 
Kyoto, B, 31: 125-197. 
MiTUi, K. 1968. Chromosomes and speciation in ferns. Sci. Rep. Tokyo 

Kyoiku Daigaku, B, 13: 285-333. 

Amherst College, Amherst, MA 01002. 

Shorter Notes 

A Recent Find of Isoetes in Louisiana. — A population of 
about two dozen plants of Isoetes melanopoda Gay & Dur. has 
been discovered in Calcasieu Parish, Louisiana. This is the first 
collection of Isoetes in the state since the 1939 collections of Brown 
and Lenz, and only the fourth collection of the species in Louisiana. 
Brown and Correll^ reported three sites, one each in Avoyelles, 
East Baton Bouge, and Rapides Parishes. The nearest to ours is 
in Rapides Parish, approximately 75 miles northeast of Lake 
Charles. CorrelP reported /. melanopoda f. pallida (Engelm.) 
Fern, in Harris County, Texas, approximately 150 miles west of 
Lake Charles. 

The plants we found grew with sedges in the marginal zone of a 
depression in a meadow. The plants have sporophylls about 20-30 
cm long, and all the collected specimens were fertile. These speci- 
mens have pale leaf bases, rather than the dark brown to blackish 
brown leaf bases of the typical form. Voucher specimens {J. Brooks 
& R. Maples 817) have been deposited at McXeese State Univer- 
sity and the University of Southwestern Louisiana. — James H. 
Brooks and Robert S. AL\ples, Jr., McXeese State University, 
Lake Charles, LA 70601^ 

^ Brown, C. A. and D. S. Correll. 1942. Ferns and Fern Allies of Louisiana. 
L.S.U. Press, Baton Rouge. 

2 CORRELL, D- S. 1956. Ferns and Fern Allies of Texas. Texas Res. Founda- 
tion, Renner. 

Shoeter Notes 187 

Jamesonia alstonii in Oaxaca, Mexico.' — The excellent 
monograph by A. ¥. Tryon,^ treating all of the species oiJamesonia, 
gives the northernmost station for the species J. alstonii and for 
the genus as Volcan Tacana, which is on the border between 
Guatemala and the state of Chiapas in Mexico. While on a bryo- 
logioal foray into the Sierra Juarez, the authors encountered 
among the grasses and herbs under scattered trees of Pimis rudis 
a fern which was not immediately recognized. Eventually the 
senior author realized that it was a Jamesonia. He tentatively 
identified it as J. alstonii, an identification kindly confirmed by 
Dr. John T. .Mickel of The New York Botanical Garden. The fern 
was collected in the highest gap on the road bc^tween Oaxaca city 
and Tuxtepec, Oaxaca, at ca 9000 ft altitude, on 26 December 
1970 {Sharp & Webster 720 A, NY, TENN). 

The occurrence of this fern at this station is of interest not only 
because it represents an extension of its range well into :\Iexico, 
but also because this is a locality from which bryophytcs of geo- 
graphical significance have been collected. Gymnomitrium lace- 
ratum, originally described from Japan but also known from the 
Great Smoky Mountains, Peru, and southeastern Africa, was 
reported there by Sharp and Hattori.^ Conostomum pentasfichum 
was also reported from the same station by Bowers and Sharp;* 
except for Mexico and Costa Rica, it seems to be restricted to the 
southern hemisphere. The senior author also has collected the 
rather rare Guamatela tuerckheimii Donn. Smith (Rosaceae), 
originally described from Guatemala, below this gap on the 
eastern slope. It seems that this area is of great significance in 
providing valual)le phytogeographical data.— A. J. Sharp and 

)J Botany, University of 

Knoxville, TX 37910. 

» Contribution from the Botanical Laboratory, University of Tennessee, 
N.S. ;i6.1, and from the Itasca Biology Session, University of Mmnesota. 
i^Tryon, Ahce F. 1962. A monograph of the fern genus Jamesonia. (_ontr. 

Gray Herb. 191: 109-197,/. -?-m ,^ .^ . , ,„„, 

3 Sharp, A. J. & S. Hattori. 1967. The range of Gymnomitrium laceratum 

extended to Me.xico. Bryol. Lich. 4: 105-106. ^ . , . p,„ .„ 

* Bowens F. D. & A. J. Sharp. 1968. Conostomum pentastichum in Losta 
Rica and Mexico. Rev. Bryol. Lich. 36: 243-244. 


American Ferx Journal 

Recent Fern Literature 

A jMonograph of the Fern Genus Eriosorus, by Alice F. 
Tryon. Contr. Gray Herb. 200: 54-174. 1970.— Alice Tryon has 
produced a model monograph of EriosoniSj the result of many 
years of painstaking work. She has studied the cytology, the 
gametophytes, the anatomy and morphology in detail; she has 
studied the plants in the field in Costa Rica, Brazil, Puerto Rico, 
Columbia, and Peru; in addition she has visited Europe several 
times and has seen most of the historic types. The taxonomic 
work can therefore be considered definitive. Although there are a 
few novelties described the general result has been to reduce many 
of the described species to synonymy. The perhaps controversial 
part of the work is the recognition of several plants as hybrids 
with the genus Jamesonia. This is reasonable in the case of 
Eriosorus elongaiiLS (Grev. & Hook.) CopeL, which is considered a 
hybrid between E, cheilanlhoides and Jamesonia, These plants 
do grow together in Andean paramos, and E. elongatus does appear 
somewhat intermediate in morphology. The matter is not so clear 
in the hybrids indicated as E, hispichdus X Jamesonia, which 
seem to show little Jamesonia influence. Some other doubtful 
plants are mentioned as variants, and so it seems that even in the 
case of a definitive monograph there will always remain some 
unanswered questions.— C.V.M. 

Index to Volume 61 

Acrostichum, 19, 41, 97-101. 119; aureum, 
97, 98, 101; calomelanos, 162; ebeneum. 
162-164; fendleri, 19; serrulatum, 117; 
speciosum, 101 

Actiniopteridaceae, 74 

Adiantum, 71, 90, 92, 174-181; capillns- 
veneris. 43, 57, 88, 90; cuneatum, 95; peda- 
tum, 45, 88, 89, 172-174; raddianum, 95 

Alsophila, 142. 143; nesiotica, 166: notabilis 

The American Species of Plagiogyria sect. 

Carinatae, 110 
Anaxetum, 37 
Anemia makrinii, 137 

Antrophyrim, 41 

An Appendageless Psilotum. Introduction to 
Aerial Shoot MorplioloKy, 75 

Aspidiaceae, 42 

Aspidium augescens, 29; blumei, 145, 148; 
germanii, 28; Krammatosorus, 148; inac- 
rounuii, 26; molle, 24; stipulare, 26; 
violascens, 24 

Aspleniopsis, 42 

Aaplenium, 43; adiantoides, 43, var. tripin- 
natifidum, 44; dayi, 162; fi lix-femina, 140; 
kamtchatkanum, 140, 141; kamtchaticum, 
141; pinnatifidum, 32-34, 45, 47; pinnati- 
fidum X trichomanes, 32, 33; platyneuron, 

Index to Volume 61 


45. 88, 89; polyodon, 43; radicans, 161; 

robustum, 44; trichomanes, 32-34, 45, 47. 


Asplenium kamtchatkaniim Gill'ert, a Dubi- 
ous Kastern Asiatic Fern, 140 

Asplenium pinnatifidum X trichomanes-A 
New Record for Indiana, 32 

Athyriaceae, 42 

Athyriopsis, 140 

Athyriuin, 140; filix-femina. 88, 140, var. 

asplenioides, 45; pterorachis, 140, 141 
Atkinson, Lenette K. Ttie Gametophyte of 

Thel\-pteris erubescens, 183 
Azolla, 1-3, 9, 11; africana, 1, 7; guineensis, 

7; irnbricata, 1, 9, var. pinnata, 7, 9, U, 

12; pinnata, 1, 3, 7-9, 12, var. africana, 7, 

A-ar. irnbricata, 7, 9, 11, 12 
Azollaceae, 42 
Belvisia. 43; mucronata, 43 
Bleclmaceae, 74, 119 
Blechnum 110; confugum, 42; diversifolium 

var. paleaceo-setositm, 44; irregulare, 44; 

tirbanii, 118; vieillartlii var. simplex, 43 
Bolbitis, 17; nmcrophylla, 18 
Botrichium dissectum f. oblitjuum, 45; 

virginianurn, 45, 88, 89 
Boutin, F. C. A New Locality for Psilotum 

nudum in Sonora, Mexico, 141 
Brooks, J. H. & R. S. Maples, Jr. A Recent 

Find of Isoetes in Louisiana, 186 
Brownlie, G. Flore de la Nouvelle-Caledonie 

et D^pendances, no. 3, Pt^ridodophytes 

(rev.), 41 
Bulbous Adder's-tongue Common in Louisi- 
ana, 39 

Camptosorus rhizophyllus, 45, 47, 88, 89 
Cephalomanes australicum, 44 
Cheilanthes fendleri, 177; lanoga, 47, 88; 
seti^era, 43 

Cibotium, 51, 55 

Cnemidaria, 142, 143 

Comparative Habitat Requirements for 

Spore Germination and Prothallial Growth 

of Three Ferns in Southeastern Michigan, 

Ctenitis, 119 

Cyathea, 142. 143, 168, 170; albifrona var. 
lata, 44; alfonsiana. 166-168; costaricensis, 
170; hastulata, 170; hemiotis, 170; holdrid- 
geana, 168-170; multiflora, 170; notabilis, 
166, 168; propinqua, 43; subg. Sphaerop- 
teris, 43; vieillardii, 43 

Cyatheaceae, 96, 142, 166, 168 

Cyclophorus, 37; americanus, 39, 94; cune- 
atus, 94;longifoHus, 39 

Cyclosorus, 26 

Cyrtomium faicatum, 44, 88, 89 

Cystopteris bulbifera, 47; fragilis, 45 
Davallia, 96; valdiviana, 96 
Dennstaedtia punctilobula, 45 
Dicksoniaceae, 166 
Diplazium, 147; acrostichoides, 45; ambigu- 

um, 161; ecliinatum, 42; expansum, 161; 

pycnocarpon, 45; radicans, 161; striatum, 

Dryoalhyriuni, 140; pterorachis, 141 
Dryopteris. 124, 127, 183; anceps, 19; 
eampj'loptera, 124, 126; campvloptera X 
intermedia, 126; celsa. 124-126; celsa X 
clintoniana. 126; celsa X {roldiana, 126; 
celsa X marginalis, 125, 126; clintoniana, 
124, 126; cHntoniana X cristata, 125, 126; 
cristata, 124, 126; cristata X intermedia, 
126; cristata X niarginalis, 126, 127; 
cristata X spinulosa. 126; subg. Cyclosorug. 
21; dentata, 24, var. harperi, 30, var. hnd- 
heimeri, 30; dilatata, 124; erubescens, 183; 
fendleri. 19; lilix-map, 124; filix-mas X 
marginalis, 126; fragrans. 1 24; goldiana, 47, 
124, 126; goldiana X marginalis, 126; 
hexagonoptera, 88, 89; intermedia. 124; 
126; ludoviciana, 124; macrophylla, 17, 
marginalis, 45, 88. 89, 124, 126; subg, 
Meniscium, 17; mollis, 24; nornialis, 27, 
28; patens var. glandulosa, 27; spinuiosa 
88, 124, var. intermedia, 45 

A Dwarf Ostrich Fern, 35 

Elaphoglossum pusilhim, 162 

Equisetaceae, 74 

Equisetum hyemale, 45 

Eriosorus, 188; cheilanthoides, 188; elongatus 
188; hispidulus, 188. var. dominicfn,sis, 162 

Tlie Fern Collections in Some European 
Herbaria-VII, 59 

The Fern Vegetation of Aldabra Atoll, 97 

Fosl>erg, F. R. The Fern Vegetation of 
Aldabra Atoll, 97 

Fox, D. L. & J. R. Wells, Schemoc}iromic 
Blue Leaf-surfaces of Selaginella, 137 

The Gametoplivte of Thelypteris erubescens 

The Gametophytes of Natural Hybrids in the 

Fern Genus Peilaea, 128 
Gastony, G. J. Asplenhun pinnatifidum X 

trichonianes-A New Record for Indiana, 32 
The Genus Grammatosorug, 145 
The Genus Selaginella in Baja California, 

Mexico, 149 
The Genug Stenochlaena J. Smith Avith 

Description of a New Species. 119 
Gibson, H. L. A Dwarf Ostrich Fern, 35 

Glaphyropteris, 183 

GlaphyropteridoPriis, 183 

Gleiclieniaceae, 73 

G6me^, L. D. Two New Tree Ferns from 

Costa Rica, 166 
Grammitidaceae, 41 
Grammatosorus. 147, 148; blumeanus, 145, 

146, 148 
Gymnogramma calomelanos. 164 

llemitelia. 142; hoidridgei, 168 

Heteroneuron meniscioides, 17 

Heusser, C. J. Pollen and Spores of Chile 

(rev.), 95 


American Fern Journal 

IlilJ, R, H. Comparative Habitat Rerjuire- 
ments for Spore Germination and Prothal- 
lial Growth of Three Ferns in Southeastern 
Michigan, 171 

Hills. L. V. (see A. Sweet) 

Holttum, R. E. The Genus Stenochiaena J. 
Smith with Description of a New Species, 

Hoshizaki,I,iarhara Joe, The Genus Adiantum 

in Cultivation [Polypodiaceae] (rev.), 95 

Howell, J. T. The Ferns and Fern Allies of 

the Sierra NevadainCaUforniaand Nevada 
(rev,), 44 

Hymenolepis, 43; revoluta var. planiuscula, 

Hymenophyllaceae, 73, 74 

Hymenophyllum le-ratii, 43; mnioides, 44, 

f. amplior, 44 
l8o6tes, 186; melanopoda, 186, f. pallida, 186 
Jamesonia, 187, 188; alstonii, 187 
Jamesonia alstonii in Oaxaca, Mexico, 187 
Lastrea erubescens, 183 
Lastreopsia, 42; tenera. 42 
Leaf Epidermal Studies in Marsilea, 101 
Lellinger, D. B. The American Species of 

Plagiogyriasect. Carinatae, 110 
Lellinger, D. B. (see C. V. Morton) 
Leptochilug fendleri, 19 
Lindsaea, 51, 53; ensifolia, 49, 51, 53-58, 

subsp. asatii, 55, subsp. ensifolia, 55; 

subg. Lindsaea, 57; odorata, 14, 16, var, 

darjeelingensis. 14-16. var. odorata, 16, 

var. pallena. 1 6; repens, 55. 57; sect. 

Schizoloma, 57 

Lindsaea (Schizoloma) ensifolia Swartz in 
Hawaii, 49 

Lindsaeaceae, 42. 74 

Lomaria adnata. 111; areolaris, 122; arguta, 
115; attenuata f. monstrosa, 44; euphlebia, 
111; fialhoi, 116; juglandifolia, 121; lenor- 
mandii f. aprica, 44; petiolata, 115; pycno- 
phylla, HI; tenuifolia, 120 

Lomaridium semicordatum, 113 

Lomariopsis, 119 

Lommasson, R. C. & C- H. Young, Jr. 
Vascularization of Fern Leaves, 87 

Lophosoria, 143 

Lunathyrium. 140; sect. Athyriopsia, 140; 

boryanum, 140; sect. Dryoathyrium, 140; 

iaponicum, 140; sect. Lunathyrium, 140; 

pterorachis. 141 
Lycopodium, 64, 137; lepidophyllum, 159; 

nutans, 43; pallescens, 160; proliferum, 43; 

aquarrosum var. pacificum, 44 
Macrothelypteris torresiana, 43 
Maples, R. S., Jr. (see J. H. Brooks) 
Marattia rolandi-principis, 43, var. sub- 

bipinnata, 43 
Marattiaceae, 73 
Marsilea, 101. 106, 109; berteroi, 104. 107; 

fournieri, 104, 107, 108; macropoda. 104. 

107; mexicana, 102. 103. 105, 107; minuta, 

108; mollis, 104, 107, 108; mucronata, 103. 
105-107; nashii, 102. 108; oligospora, 103, 
105, 106. 108; polycarpa, 102, 103, 105, 
108; quadrifoHa. 102-105, 108; tenuifolia, 
103, 104, 108; uncinata, 104. 108; vestita, 
103, 105-108 
Marsileaceae, 74 

Matteuccia, 37; struthiopteria, 36, var. 
pensyh anica, 35 

Meniacium, 17, 18; fendleri, 19; macrophyl- 

lum, 17, 18; oligophyllum. 19; sorbifolium, 

Metaxya, 143 

Mickel, J. T. & F. V. Votava. Leaf Epidermal 
Studies in Marsilea, 101 

Microlepia, 42; speluncae. 42 

Morton, C- V. Asplenium kamtchatkanum 
Gilbert, a Dubious Eastern Asiatic Fern, 
140; The Fern Collections in Some Euro- 
pean Herbaria-Vll, 59; The Genus Gram- 
matosorus. 145; The Proper Disposition of 
Meniacium macrophyllum Kunze. 17; A 
Wrongly Localized Species of Pyrrosia, 94 

Morton, C- V. & D. B. Lellinger. Niphidium 
longifolium, a Necessary New Combina- 
tion, 37; Notes on the Ferns of Dominica 
and St. Vincent-II, 161 

Nephqlea, 142, 143 

Nephrodium, 183; albescens, 26; kunthii.28; 

macrophyllum, 17 
Nephrolepis, 51, 55; biserrata, 97; cordifolia, 

43; exaitata, 51, 52, 88, 89; floccigera, 43 
A New Locality for Psilotum nudum in 

Sonora, Mexico, 141 
A New Variety of Lindsaea odorata Koxb. 

from Darjeeling in the Himalayas, 14 
Niphidium, 37, 94; americanum» 37, 39, 

longifolium, 37. 38 
Niphidium longifolium, a Necessary New 

Combination, 37 
Niphobolus, 94; americanus, 39; cuneatua, 94 
Notes on the Ferns of Dominica and St. 

Vincent-II, 161 

Notholaena cochisensis, 177. 181; grayi, 181 

Odontoloma, 57 

Onoclea sengibilis, 176, 179 

Ophiogloasaceae, 73 

Ophioglossum crotalophoroidea, 40, 41; nudi- 

caule, 51. 57, 58 
Osmunda cinnamomea, 88, 89 
Osmundaceae, 73 

Parathyrium, 140; pterorachis, 141 

Parkeriaceae, 74 

Pellaea. 128, 129, 133; andromedaefoHa, 135; 
atropurpurea, 88, 135; brldgesli, 128, 133; 
bridgesii X mucronata, 130, 132; longi- 
mucronata, 133; longimucronata X wright- 
iana, 134; mucronata, 128. 133; tenuifolia 
133; wrightiana, 133 

Pessopteris, 39 

Phegopteris, 183; macrophylla. 17 

Phlebodium aureum, 88-90 



Pilularia, 181; amerieana, 181 

Pityrop:ramma calomelanos, 162, 163; ebenea, 
162; tartarea, 162. 163 

Plagiogyria, 110, 111, 118; adnata, 111; 
subseot. Adnatae, 111; aequidentata, 115; 
anisodonta, 116; axjruta, 115; biserrata, 
113, 115; sect. Carinatae, 110-112; costari- 
censis, 113, 114; denticuiata, 116; euph- 
lebia. 111; subsect. Euphlebiae, 111; sect. 
Euplagiogyrae, 111; fialhoi, 113, 1 16; 
latifolia, 116; maxonii, 113, 117; obtusa, 
114; pectinata, 113-115; pycnophylla, HI, 
subsect. Pycnophyllae, 111; semicordata 
111. 113-115, 117; serrulata, 113, 118; 
urbanii, 118 

Plowden, C. C A Manual of Plant Names 

(rev.), 144 
Poecilopteris meniscioides, 17 
Polypodiaceae, 41 

Polypodium, 96, 183; americanum, 39; den- 

tatuni, 23; longifoUum, 39; molle, 24; 

paliiatre, 121; patens, 26; polypodioides, 

45, 47; recedens, 42; spinosum, 165; 

viri^inianuin, 45, 88; vulgare, 88, 89 
Polystichum acrostichoides, 45, 88 
Pray, T. R. The Garaetophytes of Natural 

Hybrids in the Fern Genus Pellaea, 128 
The Proper Disposition of Meniscium macro- 

phyljum Kunze, 17 
Psilotaceae. 42, 74, 83 
Psilotum, 75-77, 81-85, 141; complanatum 

81, 82; nudum, 75, 82, 141 
Pteridaceae, 42 

Pteridium, 171; aquilinum, 171, 179, var. 
latiusculum, 45 

Pteris, 119, 144; aculeata. 164; altissima, 

164, 165; arborea, 164, 165; brevinervis, 

164; bulbifera, 104; crassipes, 164; cretica, 

88, 89; data, 164; hartiana, 165; kunzeana, 

164; iactuca, 164; longibrachiata, 164; 

multisenalis, 164; regia, 164; vittata, 49, 
88, 89 

Pyrrosia, 37, 39, 94; acrostichoides. 39; 

longifolia, 39; rupestris, 94; varia, 44 
A Recent Find of Isoetes in Louisiana, 186 
Report of the 1970 Fern Foray, 44 
Reviews: The Classification of the Cyathea- 
ceae, 142; The Ferns and Fern Allies of the 
Sierra Nevada in California and Nevada, 
44; Flora de Venezuela, VoL I. Helechos, 
96; Flore de la Nouvelle-CalMonie et 
D^pendances, no. 3, Pt^ridophytes, 41; 
The Genus Adiantum in Cultivation 
[Polypodiaceael, 95; A Manual of Plant 
Names. 144; A Monograph of the Fern 
Genus Eriosorus, 188; Pollen and Spores 
of Chile, 95; 
Rouffa, A. S. An Appendageless Psilotum. 

Introduction to Aerial Shoot Morphology, 

Rumohra adiantiformis, 96 
Salvinia imbricata, 9 

Schemochromic Blue Leaf-surfaces of Selagi- 
nella, 137 

Schizaeaceae, 73, 74 

Schizoloma, 53; ensifolium. 53 

Selaginelia, 137, 138, 149, 153, 154; ser. 
Arenicolae, 149; arizonica, 150, 151» 153; 
asprella, 149, 153; bigelovii, 149. 153, 154; 
ser. Circinatae, 149, 151; cinerascens, 149- 
151, 154, 157; cugpidata, 160; eremophila, 
150. 151, 157, 158; ser. Eremophiiae, 151; 
lepidophylla, 150, 151, 157, 159, 160; 
pallescens. 150, 151. 158, 160; ser. Rupes- 
tres, 149; ser. Sartorii, 151; sul^g. Selagi- 
nelia, 149; subg. Stachygynandrum. 151; 

sect. Tetragonostachys. 149; willdenovii. 

Selaginellaceae, 74 

Sen, T. & U. Sen, A New Variety of Lindsaea 

odorata Roxb. from Darjeeling in the 

Himalayas, 14 

Sen, U. (see T. Sen) 

Sharp, A. J. & H. J. Webster, Jamesonia 

alstonii in Oaxaca, Mexico, 187 
Smith. A. R. The Thelypteris normalis 

Complex in the Southeastern United 

States, 21 
Sphaeropteris, 142 

Sphenomeris chinensis, 55, 57; chusana, 57 
Stenochlaena, 119. 122; areolaris, 120, 122; 

cumingii. 120, 122; fraxinifolia, 121; 

juglandifolia, 119, 121; laurifolia, 121, 

122; mildbraedii, 120, 121;milnei, 120-122; 

palustris, 120-122; tenuifoiia. 120 
Stromatopteris, 83; moniliformis, 83 
A Study of AzoUa pinnata R. Brown, 1 

Sweet, A. & L. V. Hills, A Study of AzoIIa 
pinnata R. Brown, 1 

Syngramma, 41 

Tectaria, 119. 145. 147. 148; blumeana, 148; 
dolichosora, 147, 148; irregularis, 148; 
melanocaulon, 147 

Teratophyllum, 119 

Thelypteridaceae, 183 

Thelypteris, 21, 43, 174-180; augescens, 28- 
31;augescens X normaUs, 31; augescens X 
ovata, 31; chrj'sodioides, 20, var. chryso- 
dioides, 20, var. goyazensis, 20; sect. 
Cj'closorus, 21; dentata, 22, 26, 31; erubes- 
cens, 183-185; gongylodes, 21; guyanen- 
sis, 19, 20; hexagonoptera, 45; X lind- 
heimeri, 30; macilenta, 25, 26; macrophylla, 
17-20; macrorhizoma, 27, 28; subg. Men- 
iscium, 17; normalis, 21, 23, 25-31; normalis 
X ovata, 31; ovata, 27-30, var. ovata. 23, 
28, 29, 31, var. Undheimeri, 23, 30, 31; 
palustris, 172, 174; patens, 26, 28. var. 
patens. 22, 26; puberula, 29; quadrangu- 
laris, 24, 25, 28 var. quadrangularis, 25, 
26, 28, var. versicolor, 22, 23, 31; reducta, 
24; saxatilis, 27, 28; serra, 29; setigera, 43; 
torresiana. 43; totta, 21, var. hirsuta, 22, 


A^MERiCAN Fern Journal 

var. totta, 22; uliginosa, 43; unca, 27, 28; 

X versicolor, 25. 26 
The Thelypteris normalis Complex in the 

Southeastern United States, 21 
Thomas, R. D, Bulbous Adder's-tongue 

Common in Louisiana, 39 
Tmesipteris, 82; tannensis, 83 
Trichipteris. 142, 168; nesiotica. 166, 168 
Trichomanea boschianum, 88, 89 
Tryon. A. F. A Monograph of the Fern Genus 

Eriosorus (rev.), 188 
Tryon, R. M,, Jr. The Classification of the 

Cyatheaceae (rev.), 142 
Two New Tree Ferns from Coata Rica, 166 
Vareschi, V, Flora de Venezuela, Vol. I. 

Ilelechos (rev.), 96 
The Variation in Spore Size and Germination 

in Dryoptehs Taxa, 123 
Vascularization of Fern Leaves, 87 

Vittariaceae, 41, 42 

Votava, F. V, (see J. T. Mickel) 

Wagner, W. H.. Jr, Lindsaea (Schizoloma) 
ensifolia Swartz in Hawaii, 49 

Wagner. W. H.. Jr. (see D. P. Whittier) 

Webster. H. J. (see A. J. Sharp) 

Wells, J. R. (see D. L. Fox) 

Whittier, D. P. & W. H. Wagner, Jr. The 
Variation in Spore Size and Germination 

in Dryopteris Taxa, 123 
Wiggins, I. L. The Genus Selaginella in Baja 

California, Mexico, 149 
Woodsia, 90; obtusa, 32, 47, 88, 89; scopulina, 


Woodwardia, 174-180; virginica, 172, 174 
A Wrongly Localized Species of Pyrrosia, 

Young, C. H., Jr, (see R. C. Lommasson) 


Page 30, line 14: For ^'Lindhemer'^ read ^^Lindheimeriy 
Page 46, line 1: For "Montgonery" read Montgomery. 
Page 53, line 9: For ^^ensifolia" read ^^ensiJoUumJ^ 


Exotic and Haray Ferns 



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In accordance with the rules and regulations of the United States Post 
Office, as established under the act of October 23, 1962, Section 4369, Title 
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Title: American Fern Journal 
Frequency of Issue: Quarterly (Approximately March 31, June 30, 

September 30, and December 31) 
Location of Office of Publication (Printers) : 3110 Elm Avenue, 

Baltimore, Maryland 21211 
Location of Business Office of Publishers (Not Printers): Dr, LeRoy 

K. Henry, Section of Plants, C^negie Museum, Pittsburg, 

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Publisher: American Fern Societt, Inc., Section of Plants, Gamete 

Museum, Pittsburg, Pa. 15213 

Editor: Dr. David B. Lellinger, Department of Botany, Smith- 
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Owner: American Fern Societt, Inc., Section of Plants, Carnegie 

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